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‘3 Volume 28 1974 

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Number 1 

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Published quarterly by THE LEPIDOPTERISTS’ SOCIETY 



\UBR aes 7 

14 March 1974 


Harry K. CLencu (Pittsburgh, Penn.) President 

ANDRE BLANCHARD (Houston, Texas) President-elect 
RonaLp W. Hopncers (Washington, D.C.) 1st Vice President 
J. C. E. Riorre (Toronto, Ontario) Vice President 

L. Vari (Pretoria, South Africa) Vice President 

S. S. NicoLay (Virginia Beach, Va.) Treasurer 

LEE D. Mumter (Sarasota, Florida) Secretary 

Members at large (three year term): R. O. Kenpauu (San Antonio, Tex.) 1975 
J. M. Burns (Cambridge, Mass.) 1974 J. A. Powe. (Berkeley, Calif.) 1975 
R. H. Carcasson (Vancouver, B.C.) 1974 J. T. Brewer (Auburndale, Mass.) 1976 
M. C. Nievsen (Lansing, Mich.) 1974 K. S. Brown (Rio de Janeiro, Brazil) 1976 
D. C. Fercuson (Washington, D.C.) 1975 K. W. Pamir (Fairbanks, Alaska) 1976 

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Memoirs of the Lepidopterists’ Society, No. 1 (Feb. 1964) 

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Tue LepiporprTreERIsts’ SOCIETY 

Volume 28 1974 Number 1 


Doucias C. FERGUSON 

Systematic Entomology Laboratory, ARS, USDA, c/o U.S. National 
Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560 

While on a field trip in Texas in June 1972, I became curious as to 
why all of the examples of Holomelina costata collected were males and 
all of the H. intermedia were females. A live female of intermedia from 
Junction, Kimble Co., was therefore kept for eggs, the larvae reared on 
Plantago major L., and adult progeny of both sexes subsequently obtained 
(Figs. 1-3). The reared males are costata and the females are intermedia, 
showing conclusively that these names as used refer to male and female 
of the same species. 

It is not especially surprising that this relationship has remained 
undetected for 85 years. The sexual dimorphism is extreme, males some- 
what resembling the immaculate form of Holomelina ferruginosa (WIk.), 
and females appearing as large, rather pale H. laeta (Guérin). Also, 
there are many other species of which only one sex, usually the male, is 
commonly collected. For example, in the same genus, hardly more than 
1% of the field collected specimens of H. ferruginosa and H. ostenta (Hy. 
Edw.) in collections are females. I have collected well over 100 speci- 
mens of ferruginosa in the Northeast without ever catching a female; 
whereas females of the aurantiaca and opella complexes are much more 
frequently taken. Thus it has been supposed that only males of costata 
were attracted to light, and that the males of intermedia were diurnal 
or for some other reason missed by the usual collecting procedures. 

The myth that both sexes of the two “species” had been collected was 
initiated by Edward L. Graef (1887: 42), who described opelloides 
from “1 ¢ and 1 2,” and intermedia from “1 ¢.” The type of intermedia 
is actually a female, and the types of opelloides are undoubtedly both 


males, although I am certain of having found only one of them. Similarly, 
the two types of Crocota diminutiva Graef, described on the same page, 
are males, although stated to be male and female. Clearly, Graef had 
difficulty determining the sex of his specimens. For costata Stretch 
(1885: 103) and cocciniceps Schaus (1901: 269) the sex of the types was 
not mentioned at all. Examination of all the material in the U.S. National 
Museum and in the collection of Mr. André Blanchard of Houston, com- 
prising a total of 198 males and 45 females, further verified the con- 
clusion that all “costata” are males and all “intermedia” are females. 

This discovery necessitates a rearranged synonymy. The types of five 
of the six names involved are in the U.S. National Museum and may be 
identified without difficulty. I have not seen the type of costata, but an 
example in the U.S. National Museum was compared with what was 
believed to be the type by F. H. Benjamin and is so labelled. This informa- 
tion, plus the original description and knowledge of what occurs in the 
type locality, leaves little doubt as to its identity. Holomelina fragilis 
(Strecker), based on a male from Pagosa Springs, Colorado, does not 
belong with costata but is very closely related to ferruginosa, apparently 
replacing the latter species in the Rocky Mountain region. 

The following revised synonymy should be substituted for that of the 
McDunnough check list (1938: 49): 

Holomelina costata costata (Stretch). Figs. 1-6. 

Crocota costata Stretch, 1885: 103. 

Type locality: Texas. 

Types: Number of specimens and sex not given. Collected by Belfrage and in 
collection of California Academy of Sciences, San Francisco. 

Crocota opelloides Graef, 1887: 42. 

Type locality: Texas. 

Types: Said to have been based on one male and one female, but both are now 
believed to be males. In the United States National Museum there are two males from 
the Graef collection that are probably the two original type specimens, but only one 
of these bears Graef’s type label. I hereby designate it the lectotype of opelloides 
(Fig. 4). 

Crocota intermedia Graef, 1887: 42. NEW SYNONYM. 

Type locality: Texas. 

Types: One female in the collection of the United States National Museum (not 
a male as stated in the original description). The type (Fig. 5) is slightly aberrant 
in having the outer dark border of the hindwing unusually wide, occupying the distal 
half of the wing. 

Holomelina costata parvula (Neumoegen and Dyar). Figs. 7-9. 

Crocota intermedia var. parvula Neumoegen and Dyar, 1893: 140. REVISED 

Type locality: Western Colorado. 
Types: Female holotype (Fig. 8), collected by Bruce, in U.S. National Museum. 

VoLUME 28, NUMBER 1 3 

7 9 

Figs. 1-9. Holomelina costata: (1) H. c. costata (Stretch) ¢, Junction, Kimble 
Co., Texas, reared 28 August 1972, D. C. Ferguson; (2) H. c. costata 2, reared 24 
August 1972 from same brood as specimen shown in fig. 1 (left forewing slightly 
deformed); (3) H. c. costata 2, Junction, Kimble Co., Texas, 18 June 1972, D. C. 
Ferguson. Parent of specimens shown in fig. 1-2; (4) H. c. costata ¢, lectotype of 
opelloides (Graef); (5) H. c. costata 9, holotype of intermedia (Graef); (6) H. c. 
costata 9, Mayer, Yavapai Co., Arizona, 23 July 1959, R. F. Sternitzky; (7) H. c. 
parvula @, lectotype of pallipennis (B. & McD.); (8) H. c. parvula (N. & D.) @, 
holotype; (9) H. c. parvula 2, holotype of cocciniceps Schaus. 

Holomelina cocciniceps Schaus, 1901: 269. REVISED STATUS. 

Type locality: Manitou, Colorado. 

Types: Number of specimens and sex not given, but the specimen labelled as the 
type in the U.S. National Museum is a female and probably a holotype (Fig. 9). 

Eubaphe costata pallipennis Barnes and McDunnough, 1918: 85, pl. 14, fig. 14. 

Type locality: Glenwood Springs, Colorado. 

Types: Described from an unstated number of male syntypes of which there are at 
least nine in the collection of the U.S. National Museum. No holotype was mentioned 
in the original description, but the specimens are labelled as type and paratypes. I 
hereby designate as lectotype of pallipennis the specimen labelled “type” in Mc- 
Dunnough’s handwriting (Fig. 7). This is not the example figured by Barnes and 
McDunnough, which, perhaps through some oversight, does not bear a type label. 
Their figured specimen is obviously one of the type lot, having been chosen to illustrate 
the new subspecies, and I think that it must be regarded as a paratype. Thus the 
type series consists of a lectotype and nine paralectotypes. 

Holomelina costata costata occurs in central Texas from Johnson and 
Palo Pinto counties, near Fort Worth, south at least to Uvalde Co., thence 


westward through the Big Bend and Davis Mountains region and southern 
New Mexico to Arizona, remaining quite uniform in appearance. Material 
from southern Colorado (and probably northern New Mexico) is some- 
what different, the males (pallipennis ) being larger and paler, the females 
(parvula, cocciniceps) having the dark border on the hindwing averaging 
narrower, and the dark brown outer border on the underside of the 
forewing weak or obsolete. As it may be considered desirable to con- 
tinue distinguishing the Colorado form as a subspecies, I have arranged 
the above synonymy accordingly. Costata in Texas has two or more 
broods, adults occurring in every month from late April to the end of 
September. For parvula, data available to me are inadequate. 

In our fauna, Holomelina costata appears most closely related to H. 
laeta, despite the normally immaculate males. Very rarely, the male of 
costata may have a complete, although very narrow, border on the hind- 
wing. There is such a specimen in the U.S. National Museum from 
Alpine, Texas, which indeed does resemble a large, pale laeta of the 
narrow-bordered form. A Mexican species, Holomelina semirosea (Druce), 
is peculiar in having males that look almost exactly like large females of 
costata, and females, if correctly associated, closely resembling ostenta. 
Such a species could be overlooked in Arizona, and collectors should 
examine their specimens carefully for any males that look like females of 
costata. The frenulum is perhaps the most convenient sex character in 
this group, being a single spine in the male and an equally long tuft of 
bristles in the female. However, the bristles of the female frenulum may 
present a deceptive appearance, being so closely appressed as to be 
mistaken for the solid spine of the male. 

I am indebted to Mr. André Blanchard and Dr. John G. Franclemont 
for the privilege of examining material in their collections. 


Barnes, W. & J. H. McDunNoucH. 1918. Notes and New Species. Contr. Nat. 
Hist. Lepid. Ni. Amer. 4(2)); 61—212) pls. 11—25. 

GrakEF, E. L. 1887. Some New Bombycidae. Entomol. Amer. 3: 41—43. 

McDunnoucu, J. H. 1938. Check List of the Lepidoptera of Canada and the 
United States of America, Pt. 1, Macrolepidoptera. Mem. So. Calif. Acad. Sci. 1: 

NEuUMOEGEN, B. & H. G. Dyar. 1893. Notes on Lithosiidae and Arctiidae with 
Descriptions of New Varieties.—I. Entomol. News 4: 138-143. 

Scuaus, W. 1901. Descriptions of some New Species of Heterocera. Ann. Mag. 
Nat. Hist., ser. 7, vol. 7: 265-270. 

StretcH, R. H. 1885. Descriptions of New Species of Heterocera. Entomol. Amer. 
1: 101-107. 

VoLUME 28, NUMBER l 5 



Department of Zoology, University of Massachusetts, 
Amherst, Massachusetts 01002 

This paper summarizes eight years of observing, rearing and collecting 
Cuculliinae by the author, primarily in southeastern Pennsylvania and 
southern New Jersey. These areas combined will be referred to as 
the Delaware Valley region. In addition, records from other collections 
are included. In all such cases, I have verified the determinations. A 
few records from the literature are also cited. However, the records 
in Tietz (1952) are generally ignored since these can not be readily 
checked, many are known errors, and many more are extremely dubious. 
A few of his most interesting records are mentioned. 

Genera for which new information can be provided are discussed in 
full. Records from the southern United States are presented due to the 
scarcity of records from that region. Taxonomy follows Franclemont (in 
Forbes, 1954). Foodplant records are from that source and the Canadian 
Department of Forestry (1962, here cited as CDF), and many are 
included which were previously unpublished. 

Major Regional Collecting Areas 


Auburn, Schuylkill County is a mixed hardwood area dominated by various oaks 
(Quercus spp.) with red maple (Acer rubrum), black birch (Betula lenta) and 
hickories (Carya spp.) as common associates. Hemlock (Tsuja canadensis) and 
Virginia pine (Pinus virginiana) are common and scattered pitch and white pines 
(P. rigida and P. strobus) occur. The elevation is approximately 850 ft. All records 
here are from Eric L. Quinter. 

French Creek State Park, Berks and Chester Cos., includes a variety of habitats 
such as dry ridges up to 900 ft. which are forested by oak sprouts of moderate to 
small size with an understory of blueberry (Vaccinium vacillans) and huckleberry 
(Gaylusaccia baccata) as well as wooded swamps dominated by mature red maples 
along with ash (Fraxinus sp.), elm (Ulmus sp.), with pin oak (Quercus palustris ) 
and tulip tree (Leriodendron tulipifera) as common associates, and spicebush 
(Lindera benzoin), Viburnum spp. and highbush blueberry ( Vaccinium corymbosum) 
forming the shrub layer. More mesic areas are forested with oaks, hickories, tulip 
tree and red maple with black birch and aspen (Populus grandidentata) also present. 
This area was collected with black light traps and bait traps, the latter in all 
habitats, by the author. 

Strafford, Chester Co., is a residential area with some patches of native woods. 
These are of two types: nearly pure stands of moderate to very large tulip trees, and 
stands of mixed oaks. In addition wild and planted fruit trees are present. The 
elevation is about 600 ft. I collected this site almost daily for over seven years, 
using both light and bait. 


Blue Mountain Bog, Schuylkill Co. is at 1600 ft. on a ridge covered largely by 
oaks. Around the bog itself trees include larch (Larix laricina) and paper birch 
(Betula papyrifera), neither of which grows generally in the region. Pitch pine 
and scrub oak (Quercus ilicifolia) occur in the immediate vicinity. Shrubs and 
sub-shrubs include cranberry and several species of blueberries (Vaccinium spp.). 
Eric Quinter has been the sole collectar at this site, using mostly black light. 


Lebanon, Hunterdon Co., is very similar in flora and moth fauna to French Creek 
(above). Joseph Muller has collected here extensively for about twenty years. This 
is referred to as Stanton by Muller (1965). 

The Pine Barrens, a unique faunal and floral region, covers much of Burlington 
Co., practically all of Ocean Co. and southward, except along the Delaware River and 
immediate coast, into Cape May County. The dry sandy “uplands” are forested 
almost exclusively with various mixtures of oaks and pines with an Ercicaceous 
understory. The exact composition is determined largely by frequent and often 
extensive forest fires that sweep through these areas, usually in April. The oaks are 
chiefly Quercus ilicifolia, marilandica, stellata, velutina and coccinea but others 
occur. The pines are almost exclusively shortleaf (P. echinata) and pitch (P. rigida) 
with stands of P. virginiana on the western fringe of the region and P. serotina and 
taeda on the southern fringe. Throughout the region are many swamps. Some are 
nearly pure stands of Chamaecyparis thyoides, known locally as cedar swamps. 
Others are mixtures of red maple, sour gum (Nyssa sylvatica) and Magnolia vir- 
giniana. In addition “pitch pine lowlands” composed of pitch pine and various 
combinations of swamp species are abundant. In all lowlands the understory is 
primarily Ericaceous. Bogs and boggy meadows are very widespread. Collectors 
interested in exploring this region should consult McCormick (1970) for more 
details on the flora. The diversity of the area as well as its combination of char- 
acteristically boreal and southern plants and moths makes it one of the truly out- 
standing collecting areas in the eastern states. 

Collecting in the Pine Barrens has been concentrated at Lakehurst where Frederick 
Lemmer, Otto Bucholz, John W. Cadbury III, and Joseph Muller have collected 
extensively. Excessive fires and cutting, and to some extent development, have 
greatly depleted the Lakehurst area, especially the swamps. Formerly all Pine Barren 
habitats were present and well collected. Other collecting sites in the Barrens 
have been at New Lisbon in a mixed swamp where Cadbury and I have collected, 
and a similar area at Whitesbog where Cadbury collected. I presently collect 
rather extensively around Batsto in a variety of habitats. The fact that all of these 
collectors have made extensive use of bait has resulted in the accumulation of a 
vast number of specimens of Cuculliinae. 

The Atlantic coastal plain outside of the Pine Barrens in New Jersey has been 
virtually uncollected for moths. Cadbury did collect at street lights in Moorestown 
and Mt. Holly, but took few Cuculiinae, as would be expected with this method. 


Members of this tribe fly during the cooler seasons only, aestivating 
as larvae in the soil during the summer. In the Delaware Valley region, 
most of the species seem to emerge in late October or November, although 
in 1971 they did not appear in numbers until well into December. 
October had been very warm. However, Metaxaglaea viatica adults 
emerging in early October 1972 remained lethargic until early November 

VoLUME 28, NuMBER 1 i 

when mating occurred. There is strong circumstantial evidence that 
other species may be present but inactive in October. Most of the 
species overwinter as adults, at least sometimes in the fallen leaves on 
the ground. In these species development of the eggs within the females 
is not evident upon dissection until at least the end of January. Mating 
occurs from late January to mid-March in Eupsilia spp., by mid-March 
in Lithophane grotei, but apparently usually not until well into April in 
Lithophane hemina, patefacta, querquera, and viridipallens. The fact 
that egg development occurs in at least some species during midwinter 
seems to indicate that no true diapause occurs. In addition large num- 
bers of Eupsilia spp. can be taken at bait on almost any warm winter 
night, when the temperature is above 42°F. Lithophane spp. are also 
taken on such nights, but not in large numbers. Rainy nights are generally 
far more productive than clear ones when the temperature is lower than 

Even when nights are too cold for flight, the moths are not completely 
inactive. Caged Eupsilia and Lithophane will crawl out of their shelters 
to take water on rainy afternoons, and also will crawl about probing 
with their proboscis on sunny winter days. In either case the temperature 
is often under 40°F. On cold nights the moths may crawl about in search 
of better shelters, which are located at least in part by probing with 
the antennae. This has been observed repeatedly at subfreezing temper- 
atures for a variety of species: Lithophane viridipallens, antennata, 
unimoda, grotei, and patefacta; Eupsilia vinulenta, sidus, morrisoni; 
Epiglaea decliva. The minimum temperature for activity seems to 
per 23or 

Suitable shelters are essential for winter survival. The moths always 
rest with the abdomen pressed against some surface and the wings closed 
tightly over it. The thorax is not covered, but is densely hairy. The 
head is tucked up tight against the thorax and the antennae are folded 
under the wings. One night, three Eupsilia vinulenta were dislodged 
in a cage and fell onto their backs. The temperature was about 28°F. 
By morning, two were dead, but the other had righted itself and crawled 
under a leaf. About a dozen others in the box, under leaves or on the 
sides, survived the night which fell to at least 21°F. For prolonged 
survival the moths crawl deep into leaf litter in cages. Almost invariably 
they crawl into a folded over leaf. Probably water retention, rather 
than direct temperature effects, necessitate the use of shelters. Re- 
frigerated moths will survive several months in tightly closed con- 
tainers, but only a few days in well ventilated ones, in frost-free 
refrigerators. Similar observations have been made outdoors in cold 


spells. In fact, it is quite possible that lack of protective snow cover in 
the southern states may be a limiting factor for these moths. Certainly, 
they are essentially a northern group. 

Little is actually known of the resting habits of these moths in the 
field. I have seen Eupsilia and Sunira bicolorago fly up from leaf litter 
on several occasions. However, T. D. Sargent (pers. comm.) has taken 
Eupsilia vinulenta at light in Massachusetts on nights when the ground 
was completely snow-covered. Presumably these moths had been under 
loose bark, or in hollow trees. The brown Lithophane look as if they 
might hide under loose bark, although caged individuals crawl under 
leaves. Certainly, Lithophane lemmeri looks like a perfect match for 
the bark of its larval host, white cedar. In addition, cedar swamps 
almost always flood in the winter. It seems that this species must over- 
winter in the shreds of bark that are typical of old cedars. Lithophane 
thaxteri and lepida also resemble the bark of some conifers, notably 
pitch and shortleaf pine. L. pexata also resembles bark in general. All 
three of these species characteristically rest on the sides of bait traps, 
head up. Other Cuculliinae rest in the dead leaves provided at the 
bottom of such traps. 

In those species that do not hibernate as adults, the egg is the over- 
wintering stage. These are laid in the autumn or early winter, well into 
December, in Metaxaglaea viatica and Epiglaea decliva. 

Larvae of this tribe hatch in the spring from late March into 
early May depending on the species. Most of them can survive for 
well over a week without food, provided humidity is high, at room tem- 
peratures. This ability would minimize losses from early emergences. 
Most or all of the species will accept catkins and these are apparently 
the normal initial food source for Sunira bicolorago and Anathix spp. 
(Forbes, 1954). I am aware of only one species, an undescribed 
Metaxaglaea, which is an obligatory catkin feeder. I find that Sunira 
larvae will eat any part, including the wood, of practically any plant. 
Unlike most of the species however, Sunira larvae are not predatory. 
In general, most species probably start feeding on newly opened leaf 
buds. As the larvae mature they feed on the leaves and flowers and, in 
some Lithophane at least, on fruits as well. In my experience Lithophane 
larvae are extremely predatory in captivity. It seems likely that they 
eat other caterpillars in natural conditions as well. This would certainly 
be an adaptive habit when competition with other species is acute as 
sometimes happens during canker worm (Geometridae) outbreaks. 
Cannibalism might also be adaptive as a means of population control. 
Lithophane larvae will also eat dead larvae and frass in captivity. 

VoLUME 28, NUMBER 1 9 

In general the moths are most often taken at a variety of baits. I usually 
use a mixture of rotten apples, crushed bananas, brown sugar, and 
molasses. This mixture is allowed to ferment for at least three days 
before use. Beer may be added if fermentation is inadequate. These 
baits last about one month, longer if they are refrigerated when not 
in use. Naturally rotted apples and quinces also make excellent bait. 
Baits may be smeared on tree trunks or placed in dishes. Apparently 
they must be at least four feet above the ground. Joseph Muller and I 
have also had success with bait traps. Ours are similar in basic design 
to that described by Platt (1969) except that they are made of metal 
with pie pans as lids. Great care must be taken to assure that the lids 
fit snugly. These traps are hung from tree limbs. I check mine once 
a week or less. 

Another excellent method of collecting these and other noctuids is 
to net them from pussy willow blossoms just after dusk. The moths 
may also be obtained by shaking the tree as they will usually fall to the 
ground rather than take flight. I have taken six species of Lithophane 
and three of Eupsilia at pussy willow at Strafford. This total includes 
all except the rarest members of the tribe that fly at that site in the 
spring. Other blossoms might prove effective as well. Holland (1903) 
notes that “Scopelosoma” (i.e. Eupsilia and Pyreferra) come to maple 
sap buckets. 

In general, lights tend to yield poor returns. However, the setup used 
by Sargent at Leverett, Massachusetts is quite effective. This consists 
of a Robinson Trap, a 15-watt black light, and four 150-watt flood- 
lights. On 16 March 1973, Sargent and I were able to compare his lights 
with ten baited trees. Twenty-two species were taken. A total of 212 
specimens of 21 species was taken at the bait, while 69 specimens of 
14 species were taken at light. During the course of the spring, bait 
proved most effective for all members of the tribe except Homoglaea 
hircina which appeared only at light. In general the proportion of speci- 
mens at light seems to increase sharply in mid-April in both Pennsylvania 
and Massachusetts. 

The effects of weather on the activity of these moths are only partially 
understood. Observations on captive individuals suggest 47°F as the 
minimum temperature at which Lithophane patefacta will take flight. 
Similarly, the minimum for Eupsilia spp. was found to be 42°F. However, 
clear cold nights following warm, sunny, spring days will often produce 
substantial catches even with temperatures at dusk as low as 36°F. 
Apparently the hairiness of the moths enables them to retain body heat, 
even with rapidly falling temperatures. Such captures are almost always 


within one hour of dusk and usually at light. Certainly, any night with 
the temperature at dusk above 50°F is likely to be productive. 

Xylena Ochsenheimer 

The larvae of this genus are brownish, climbing cutworms in the last 
instar; green with white markings in the earlier instars, resting then in 
the foliage. They appear to be very general feeders. 

X. nupera (Hubner) has been taken at Lakehurst by Muller and 
Cadbury, and once at Batsto by myself. Dates span from 16 September to 
30 April. It is clearly very rare in the region. Tietz (1952) records it 
from Delaware and Berks Cos., possibly correctly. 

X. curvimacula (Morrison) is quite general in the region. It is common 
only in Schuylkill Co. where Quinter finds it at several sites. It is very 
scarce in the Pine Barrens and not taken at Batsto. The moth flies 
from late October to late April, most captures are in April. 

X. cineritia (Grote) was taken by Lemmer a few times at Lakehurst 
in the 1930’s and 1940’s. Quinter has taken it in substantial numbers at 
Blue Mountain Bog, Auburn, and nearby New Ringold. Tietz reports 
it from Reading, Pennsylvania. The species is not known from south 
of this region. Records are from October to April. 

Lithophane Hubner 

This genus is somewhat heterogeneous. The first group, through 
oriunda, is distinctive in adult maculation and genitalia. The larvae 
are brown or gray in the last instar with a mottled pattern as described 
by Franclemont (in Forbes, 1954). The earlier instars are green with 
the ordinary lines and tubercles contrastingly white. The green instar 
larvae rest by day on the foliage of the host tree. The last instar of at 
least bethunei is a climbing cutworm, resting by day in bark crevices 
and among debris at the base of the tree. One last instar hemina larva 
has been found on a twig of boxelder. 

The generic name Grapolitha (Hubner) was based on L. socia, a 
European member of this group (Franclemont, 1942). The adults of 
most of these species are dimorphic (Franclemont, 1942; Forbes, 1954). 
Adults of this group are very rarely taken in January and February 
in this region. 

L. semiusta (Grote) is extremely rare in the region. Lemmer took one 
at Lakehurst dated 1 to 10 May (specimen in the American Museum), 
and Muller has taken one at Stanton, 18 October 1953. Lemmer did 
not include years on his labels. Tietz (1952) states that the United 
States National Museum has one from White Mills, Wayne Co., Penn- 

VoLUME 28, NUMBER 1 ill 

sylvania dated in August. The species is northern and Lakehurst is the 
most southern capture. It ranges west at least to Wausau, Marathon 
Co., Wisconsin (Jim Parkinson, in Schweitzer collection). The larva 
occurs on basswood and accepts choke cherry (Forbes, 1954; CDF). 

L. bethunei (Grote and Robinson) is fairly general in the region except 
for Schuylkill Co. where Quinter has not taken it. Otherwise it is usually 
common outside of the Pine Barrens, where however, it is present. Larva, 
on many trees (Forbes, 1954; CDF’). I have reared them on oak, Prunus 
spp. and apple, and have found two larvae on trunks of black oak trees 
in bark crevices at Springfield, Delaware Co., Pennsylvania. It flies 
from October through April, but is taken much more often in spring. Only 
the typical form has been taken locally. 

L. innominata (Smith) was taken at Lakehurst by Buchholz ( American 
Museum) from October to April, and at Auburn, 28 April 1972. Other- 
wise the only regional record is the Wayne Co., Pennsylvania specimen 
noted by Tietz (1952) as being in the United States National Museum. 
There are no records from south of this region. Only the normal form 
illecebra (Franclemont) has been found in the area. 

L. patefacta (Walker) has been taken by all collectors at all Pine 
Barren sites. It is very common some years, very rare others. It does 
not occur, so far as known, anywhere else south of the vicinity of Ithaca, 
New York. Tietz’s record (1952) for western Pennsylvania is probably 
an error, and Muller's record (1965) from Stanton is known to be so. 
I have reared it on Prunus spp. and commercial blueberry leaves. It 
eats the leaves, flowers and fruit of the former. The typical and 
niveocosta (Franclemont) forms are of nearly equal abundance. Mating 
takes place in late March and early April. 

L. hemina (Grote) is apparently general in the region, but is quite 
rare in the Pine Barrens where it has been taken only at Lakehurst by 
Cadbury and Lemmer. It is common at Lebanon, Auburn, and French 
Creek, where it seems to have no habitat preference. It is not common 
at Strafford. Other area records are Valley Forge Park, Pennsylvania, 
and Moorestown, New Jersey. It flies from mid-October to as late as 
30 April. I have attempted four times to obtain ova from females baited 
in early April, but all laid only sterile eggs. I have also found a larva on 
a boxelder (Acer negundo) twig at Wissahickon Ravine, Philadelphia, 
11 June 1973. The only known records south of this region are a specimen 
taken by Quinter at black light at Ice Mountain, Hampshire Co., West 
Virginia, 24 April 1971; and an apparent specimen from C. V. Covell 
taken at Valley Station, Kentucky, near Louisville, November 1972 by 
Siegfried Schloss. Twenty out of 32 regional specimens in the author's 


collection are the variety lignicosta (Franclemont). It feeds on many 
trees in captivity. 

L. petulca (Grote) ranges as far south as this region. It is probably 
common at Auburn since a small series has been accumulated solely at 
light. I took it once at French Creek (genitalia checked). Muller gets 
it at Lebanon. The American Museum has eight specimens under this 
name from Lakehurst, but some are almost certainly L. signosa. At least 
two are correct, however. Both forms occur in the region. Usually, 
there is some violet shading at least along the costa of the primaries. 
The larva feeds on many trees (CDF). 

L. signosa (Walker) is not common, but is found at Strafford, French 
Creek, Lebanon, Batsto, Moorestown, Lakehurst, and Whitesbog. I also 
took one in a crevice on the trunk of a ginko tree at Overbrook, Phila- 
delphia Co., Pennsylvania, 18 November 1968. The larva and food are 
unknown, but the species is apparently a restricted feeder ( Forbes, 1954). 
This is apparently the most southern species of this group. Its known 
range is from Bristol, Rhode Island and Randolph, Vermont (Francle- 
mont, 1942) to Arlington, Virginia (Forbes, 1954) and Clinton, Hinds 
Co., Mississippi (Bryant Mather, 10 March 1960; determined and geni- 
talia checked by the author, form pallidicosta Franclemont), and 
Lafayette, Indiana (Franclemont, 1942). The lack of records from 
Canada is noteworthy. Nine of my seventeen regional specimens are 
the form pallidicosta. One of these has the suffused area reddish brown 
instead of the usual blackish, as does the Mississippi specimen. Dates 
range from 11 October (Strafford) to 4 May (Moorestown, 1941). 

L. disposita (Morrison) is extremely rare in the region. It has been 
taken at Lakehurst, 20 April 1952 (Muller) and 18 October 1946 (Cad- 
bury); Wayne, Delaware Co., Pennsylvania, 16 March 1965 (Schweitzer, 
on store front); Auburn, 24 September 1972; and Philadelphia, 19 No- 
vember 1902 (Quinter coll.). Tietz reports it from White Mills, Wayne 
Co., in August, based on a specimen in the United States National 

L. oriunda (Walker) has been taken twice at Scranton (AMNBH). 

The larvae of the following species (second group) are green in all 
instars so far as known and remain in the foliage by day. The adults are 
generally shades of grey and are usually monomorphic. The adults are 
fairly frequent in mid-winter. Mating seasons are variable between 
species. Some of the larvae are restricted feeders. The name Lithophane 
was based on a species related to our pexata. 

L. lamda (Fabricius) race thaxteri (Grote) is reported in eastern 
Pennsylvania at White Mills and Reading by Tietz (1952). This species 

VoLUME 28, NUMBER 1 13 

would be hard to misidentify. I cannot verify its presence in that state. 
It is, however, common some years in the Pine Barrens where it has 
been take at Lakehurst (Cadbury, Lemmer) and Batsto. The local 
foodplant is unknown. The host reported by Forbes (1954), Ceanothus, 
is not present in the Barrens. Myrica is listed for the typical, European, 
race and Douglas fir (Pseudotsuga) for the Pacific Northwest popu- 
lation. Brower has one from Maine (ex larva) on Myrica gale but 
throughout much of Canada the principal host is larch (CDF). Thus 
our population might be expected to feed on Myrica pennsylvanica or 
pine or white cedar. It is possible that several species are involved. 
It seems certain that this is not a general feeder. It flies October to 

L. pexata (Grote) is not common in the region, but has been taken at 
Lebanon, Lakehurst, Weekstown (Atlantic Co., Pine Barrens), French 
Creek and in Schuylkill Co. at Auburn, New Ringold and Blue Moun- 
tain Bog from October to April. The principal host seems to be alder, 
but birch has also been reported (CDF). Quinter finds only the alder 
to be acceptable. 

L. fagina (Morrison) is a northern species that extends south into the 
Pine Barrens where it is usually rare. It has been taken at Batsto, New 
Lisbon, and Lakehurst. Area dates are from 21 September (Cadbury) 
to mid-April. In Massachusets, the species seems to mate rather late 
in the spring. The larva has been collected in Canada on birch twenty 
three times and once on pin cherry (CDF). Presumably, the food in 
the Pine Barrens is Betula populifolia. The young larva is similar to L. 

L. tepida (Grote) is very rare in this region, having been taken at 
Lebanon, 23 November (Muller); and a few times at Auburn by Quinter. 
It has been reported from White Mills in August (Tietz, 1952). I have 
reared the larva (from Massachusetts ), and birch and aspens seem to be 
the preferred hosts. It is very predatory, and will run down other larvae. 
The last instar is a somewhat yellowish green with slightly broken yellow 
ordinary lines and tubercles. It has been found on birch, willow, and 
gooseberry in Canada (CDF). 

L. baileyi (Grote) is reported by Tietz (1952) from White Mills, in 
August, and has been taken at Lakehurst, 16 October 1946 (Cadbury). 
The larva has been reported on pine, birch and cottonwood (CDF). 

L. querquera (Grote) is very rare in the Pine Barrens, having been 
taken only at Lakehurst by Lemmer and once at Batsto, but is less rare 
elsewhere. Records are for Lebanon (common), Auburn and French 
Creek. It may prefer red maple swamps. This species ranges south to 


Clinton, Hinds Co., Mississippi where Mather has taken it once, 20 
January 1969. I have seen the specimen. In the Delaware Valley 
region, both forms occur at all localities. The larva of this species is 
unique. It is uniform dark leaf green. The ordinary lines and tubercles 
are not visible except for a contrasting but poorly defined yellow 
spiracular line. There is a bright yellow transverse line at the posterior 
of each segment except on the anal hump. The anal hump and prolegs 
are contrastingly white. The ground color becomes gray in the last instar. 
It seems very unlikely that this is a cryptic species. It will accept a variety 
of hardwoods, but the newly hatched larvae seem to prefer paper birch 
and seem to dislike black oak. The larvae are extremely predacious and 
will also eat human epidermis when handled. 

L. viridipallens (Grote) is usually not rare in pitch pine lowland habi- 
tats in the Pine Barrens where it has been taken at all localities collected, 
although it has not been taken at Lakehurst in over twenty years. Most 
records are from November to April, but there are a few in October. 
All specimens seen by me have been from along the Atlantic and Gulf 
coasts from Hampton, New Hampshire (Shaw, in Brower coll.) to 
Carteret Co., North Carolina (J. B. Sullivan) and McClellanville, South 
Carolina (R. B. Dominick) and Lauderdale Co., Mississippi (Mather). 
Tietz’s reports (1952) from western Pennsylvania are extremely dubious. 
Nothing is known of the life history except that mating apparently does 
not take place until well into April in the Pine Barrens. 

L. lemmeri (Barnes and McDunnough) occurs in the Pine Barrens 
where Cadbury and Lemmer found it abundant some years at Lakehurst, 
as late as the 1940’s. More recently, Muller has not found the species 
and I have taken but one at Batsto, 12 April 1970. Dates are from 
October to 12 May. The food is white cedar according to Muller 
(1965). This information is from Lemmer who presumably reared it. 
Franclemont (1969) suggests that the food may be red cedar, but the 
distribution and habitat of the moth make this seem very unlikely. Other 
records are Ivoryton, Connecticut (Forbes, 1954) and McClellanville, 
South Carolina (Dominick). The South Carolina specimens seemed 
atypical, but I did not have other material available for comparison. 

L. lepida (Lintner ) is a northern species ranging into upper New York 
State in its typical form. The race adipel (Benjamin) occurs only in 
the New Jersey Pine Barrens and was sometimes taken abundantly by 
Cadbury and Lemmer at Lakehurst, and was common in December 1972 
at Batsto, but very rare other seasons. I have found it only in pitch 
pine lowland habitats, despite more extensive baiting in drier sites. The 
type race feeds on pines (Forbes, 1954; CDF). 

VoLUME 28, NUMBER l 15 

L. antennata (Walker) is completely general in this region and always 
common, especially at French Creek. It is often numerous as early as 
late September and occasionally flies into early May. The larva feeds 
on most hardwoods, eating fruits and galls as well as leaves (Holland, 
1903; Forbes, 1954). I have seen specimens from Hinds Co., Mississippi 
(Mather) and have a series from Pine Mt., Kentucky, 22 October 
1970 (Cornett, received from Covell). 

L. grotei (Riley) is another common, generally distributed species in 
the Delaware Valley region. It appears later and disappears earlier 
than antennata, most records ranging between November and March. 
I have reared it from eggs found on Prunus serotina at Strafford, and 
Brower informs me that it sometimes causes substantial defoliation of 
soft maples in Maine. The early mating and oviposition suggest that 
the larvae may begin to feed on catkins in some cases. 

L. laticinerea (Grote) is surprisingly rare in this region and most 
specimens seen labelled this are grotei. It is known from Lakehurst, 
Batsto (one, November 1968; and a larva on Quercus velutina, 25 May 
1969), Strafford (1 November and 19 December 1967, 22 January 1973 
and probably 8 April 1970), and Auburn, where it is also quite rare. 
The species ranges south to Clemson, South Carolina (22 February 
1939, E. C. Sturgis, in Schweitzer coll.). Dominick has a specimen 
from McClellanville, South Carolina which superficially resembles this 
species, taken 16 February 1970. This species closely resembles antennata 
but averages larger, and slightly darker and duller. However, at least 
in the Delaware Valley area it is best to check the male genitalia with 
the figure in Forbes (1954). The presence of a basal dash on the 
primaries will distinguish this species from grotei which is also much 
darker and more mottled. 

L. unimoda (Lintner ) is completely general in this region and is usually 
common except perhaps in Schuylkill Co., where, however, more collect- 
ing at bait might turn up larger numbers. It seems to be common in 
almost any habitat. Franclemont (in Forbes, 1954) reports that the 
larva feeds by preference on black cherry, but the moth is clearly not 
associated with this plant in this area. 

Eupsilia Hubner 
The moths of this genus make up the majority of mid-winter moth 
catches in this area. They fly from late September (very rarely) into 
late April. Eggs develop in January and matings occur from the end of 
that month into March. The moths are sometimes difficult to identify 
but the following generalizations seem safe. E. vinulenta (Grote) and 


E. sidus (Guenée) are general and common in the region and fluctuate 
in numbers from year to year. E. morrisoni (Grote) is usually rare 
in the Pine Barrens, but common elsewhere. E. tristigmata (Grote) 
is common in the Pine Barrens and general, but rare, in the rest of the 
region. E. cirripalea (Franclemont) may be common in the Pine Barrens, 
but is definitely rare elsewhere, though taken at Strafford and the 
Nottingham (Chester Co.) pine barren area. E. devia (Grote) is rare 
throughout the region, except at Lebanon. It is unknown from Schuylkill 
Co. so far. The larvae of all the species are described by Forbes (1954). 
I have reared the larvae of most of the species on various trees and shrubs. 
They will eat dandelion as well. The young larvae make a crude silk 
nest between two leaves. They are solitary. The mature larvae hide in 
debris at the base of trees and probably in bark crevices as well. 

Pyreferra Franclemont 

P. hesperidago (Guenée) has been taken at Lebanon, Auburn, Strafford 
and Valley Forge Park which are within the range of the foodplant, 
Hamamelis, reported by Forbes (1954). However, this plant does not 
grow in the Pine Barrens although the moth has been taken a number of 
times by Cadbury at Lakehurst. Possibly, sweet gum, Liquidambar 
styraciflua, which is in the Hamamelidaceae is the foodplant there. 

P. citromba (Franclemont) has been taken at the same places as the 
last species. Likewise, its reported foodplant, Corylus, does not grow in 
the Pine Barrens. Birch, Betula populifolia, seems to be the most likely 
host there. Corylus is also quite uncommon in the Delaware Valley 
region as a whole. 

P. pettiti (Grote) has been taken once by Muller at Lebanon. Since it 
feeds on Betula lenta and B. lutea according to Forbes (1954), its 
rarity in this region is remarkable. 

P. ceromatica (Grote) has not been taken in this region so far as I 
am aware. 

All of the species apparently fly from October through April. They 
have not been taken in mid-winter. 

Homoglaea Morrison 

H. hircina (Morrison) is somewhat surprisingly established this far 
south. Muller has a series from Lebanon; Quinter has several from 
Auburn; and I have one from Strafford. It flies from October to April. 
The larvae web together aspen leaves upon which they feed (Forbes, 


Sericaglaea Franclemont 

S. signata (French) is fairly common in the Pine Barrens, having been 
taken at all the usual locations, including a swamp near Weekstown and 
also at a site east of Berlin (Quinter). Otherwise, the only area records 
are singles from Strafford, and French Creek. Dates run mid-October 
into April, except the Berlin specimen which is 2 May 1970. I have 
examined the Florida specimens in the American Museum and am fairly 
sure they are variants of C. tremula. However, the species appears to be 
widespread in the south. Bryant Mather takes it commonly in Hinds Co., 
Mississippi, and Dominick has a series from McClellanville on the coast 
of South Carolina. Ric Peigler has taken it at Greenville on the Piedmont 
of that state. The Clemson University Entomology Department has one 
from Florence, also on the coast. In the south dates are December to 

The genus Metaxaglaea (Franclemont) will be treated in a later 
paper which will include a description of a new species from the Pine 
Barrens and south. 

Epiglaea Grote 

E. decliva (Grote) is completely general in this region. It is usually 
not uncommon, but the only record of it occurring in abundance is at 
the pine barren area near Nottingham, Pennsylvania, 16 October 1971, 
when each of about 35 baited trees had several individuals each time they 
were checked. Since this is the earliest regional date, it is likely that 
the species had not yet reached peak abundance. The latest record is 
for seven specimens at Strafford, 31 December 1972. The females in this 
lot produced largely sterile eggs but each produce some viable ones. I 
once found a last instar larva feeding by night on the lowest limb of an 
apple tree at Strafford. I have reared the species on Prunus spp. The 
larvae rest by day on the branches. The last instar larva probably 
hides in bark crevices or on the ground. 

E. apiata (Grote) is nearly limited to the Pine Barrens in this region. 
It is general and abundant in them from late September into November. 
The only other locality for the species is Lebanon where Muller took 
a male on 10 April 1952. This date is quite remarkable. The specimen 
appears too fresh to have overwintered. The normal food is cran- 
berry, but it has been reported from blueberry (Forbes, 1954). The 
species occurs in the coastal plain south to North Carolina (Fort Bragg, 
R. A. Anderson, Carteret Co., J. B. Sullivan) and McClellanville, South 
Carolina (Dominick). Southward, most records are in November. North- 
ward, the species is not limited to the coast. 


Chaetaglaea Franclemont 

C. sericea (Morrison) is a very widespread species and is quite general 
in the Delaware Valley region. It is common all over the Pine Barrens. 
It was also taken in numbers in a burned over area dominated by sprout 
oaks over blueberry at Resica Falls, Monroe Co., 28 October 1971, 
and on the pine barren area at Nottingham, Chester Co., 16 October 
1971. Otherwise it is uncommon in the region in my experience. The 
species occurs on the coastal plain in North Carolina (Carteret Co., 
Sullivan) and South Carolina (Florence Co., Clemson University coll.; 
McClellanville, Dominick). Kimball (1965) reports it in Florida. I 
have also seen one from Mather’s collection from Newton Co., Mississippi. 
In the Delaware Valley area, the species flies from late September into 
December, mostly in October. Forbes (1954) describes the larva and 
I have reared it on Prunus spp. 

C. tremula (Harvey) ranges along the coast from Bar Harbor, Maine 
(Brower) to Florida and Texas. Brower also has it from inland at 
Scranton, Pennsylvania and Bear Mountain, New York. In the Delaware 
Valley region it is abundant in the Pine Barrens mostly in late September 
and early October, but not taken elsewhere. I have reared it on Prunus 
spp. and find the larva identical to sericea except that some individuals 
have a blue tint dorsally. The eggs of this species are usually attached 
weakly, if at all, to any substrate and presumably fall to the ground in 
the winter. Thus the larva probably feeds on shrubs. 

C. cerata (Franclemont) has an unusual distribution, ranging from 
Mystic, Connecticut (holotype, Franclemont, 1943), up the coast to at 
least Hampton, New Hampshire (Shaw, in Brower coll.) and also inland 
at Augusta, Maine (Brower, 18 September i968). It also occurs in 
Pennsylvania at Finleyville (presumably the one in Allegheny Co., Engel’s 
usual collecting area, although there is such a town in Bedford Co. 
as well), and at Auburn where it is rather common. All records, except 
the Maine one, are in October. 

Psectraglaea Hampson 

P. carnosa (Grote) is general and often common in the Pine Barrens, 
at least south to Batsto. Adults may often be found on the red leaves 
of blueberry and huckleberry (Muller, Cadbury, Forbes (1954) ). They 
also come to bait and light. In Pennsylvania, it is known from near Mt. 
Pocono (T. Lis, in Schweitzer coll.) and reported from Drifton, Luzerne 
Co., September (Tietz, 1952). Presumably the Drifton specimen is the 
basis of Forbes’ (1954) Luzerne Co. record. Most records are in October, 
but it flies into November. Muller (1965) reports huckleberry (Gaylusac- 

VoLUME 28, NUMBER 1 19 

cia) as the foodplant. Darlington (1952) writes that Buchholz reared the 
species on wild cherry and that blueberry was unacceptable. 

Anathix Franclemont 

A. ralla (Grote and Robinson) is fairly common at French Creek, 
Auburn and Lebanon; less common at Strafford; and not seen from the 
other sites, although Rummel is reported (Anon., 1923) to have taken a 
specimen, 6 September 1922, at Lakehurst. It flies in this area from late 
August into October, most commonly in late September. 

A few genera in this tribe have not been discussed above. Of these, 
Hillia, Xanthia and Lithomoia are apparently not represented in the 
Delaware Valley. The others contain widespread species which are 
general and common in the region. 

Anytus Grote 

A. privatus (Walker) is known in this area by only one definite record, 
Resica Falls, Monroe Co., Pennsylvania, taken 25 August 1971 by the 
author. It is not different in any way from my New England series. 
Tietz (1952) reports it from Berks Co. in July. 

A. teltowa (Smith) may be conspecific with privatus (Forbes, 1954). 
If so, I would retain the name as a subspecies. It has been taken in the 
Pine Barrens numerous times in late August and September, but it is 
seldom if ever really common there. I have seen similar specimens from 
coastal North Carolina (Carteret Co., Sullivan) and coastal South Caro- 
lina (Florence, in Clemson University coll.). Quinter has taken it once 
at Auburn. 

Xylotype Hampson 

X. capax (Grote) has been taken throughout the Pine Barrens where 
it is almost always common and sometimes locally abundant. Otherwise, 
it is known from Blue Mountain Bog and Nottingham Barrens. It is 
common at both places. It is also reported from Flourtown, Montgomery 
Co., Pennsylvania by Shapiro (1965). The American Museum has two 
Rothke specimens from Scranton. In cases where the habitat is known, 
pitch pine and scrub and/or blackjack and post oak have been common 
in the immediate vicinity. The primaries would be an excellent match 
for pitch pine bark. Most records are mid-October but there are a few 
in September and November. I have reared the larvae on a diet of 
red oak, wild cherry, crabapple and blueberry leaves. Pitch pine is not 
accepted. Forbes suggests Vaccinium is the preferred foodplant, but 


species of this genus are not common at the Nottingham site. I suspect 
that the larvae feed on a variety of plants, but if they are restricted 
feeders, one of the shrubby oaks seems most likely in this area. In 
Alberta and Saskatchewan there is a spruce feeding population (CDF) 
but I suspect these may not be conspecific with capax. Larvae of X. 
acadia, which Forbes (1954) treats as a race of capax, have been reported 
from alder (3) and larch (1) (CDF). I suspect that the species may be 
limited by some habitat requirement unrelated to foodplant. Certainly, 
this is a very local species. A larval description follows: 

Last instar: Head and true legs, brownish red with darker shading. Cervical 
shield, nearly black with 12 minute white dots. Ground of body somewhat violet 
gray, strongly mottled with paler shades. Dorsal line, faint or absent. Subdorsal 
line represented by a conspicuous white dot in middle of all segments except first 
and last. Anterior to, and dorsal to these, a smaller white spot connected to sub- 
dorsal spot by a black patch, except on second and third segments. Lateral line, 
absent. Stigmatal line, very broad, white with definite blue tint, enclosing spiracles. 
Above and anterior to spiracles, a white dot with a black bar running to anterior 
edge of segment. Laterally, a conspicuous orange patch on each segment except 
first and last three. Anteriorly, ground shades into a dull, pale brown, partially 
obliterating pattern. Below stigmatal line, ground mottled with red, and with a 
white spot on most segments. Prolegs with a green patch. 

Earlier instars: Head reddish, with darker patches. Anal segments enlarged, 
forming a hump, colored as rest of body. Body purplish, ventral surface not paler; 
subspiracular line very broad, cream color. Dorsal and subdorsal lines fine, cream. 
Pair of faintly darker patches centered about dorsal on each segment. Prolegs paler 
pinkish. This description, based on third and fourth instar larva. First instar is a 
semilooper as are most Noctuidae. The larva hides under debris in the later instars, 
at least in captivity. Unlike the Lithophanini, this species pupates in June and 
diapauses in that state. 


The moths of this subfamily are primarily northern. Many do extend 
into the Delaware Valley region. Some extend much farther south. 
In general, they are less common in the southern parts of their range, 
although space did not allow for full discussion of this in the text. A 
few of the species are essentially southern. From what I have seen of 
southern collections there seems no doubt that these moths are poorly 
represented in that region and not merely overlooked. Several new life 
history data are presented here as well as new distributional data. I 
strongly suspect that more thorough collecting in the southern Appa- 
lachians will turn up many species, quite possibly some endemics. The 
Mt. Mitchell, North Carolina area seems especially Pron I hope 
that this paper will stimulate interest in these moths. 


I wish to thank the following persons for the use of their collections: 
Dr. A. E. Brower, Augusta, Maine; J. W. Cadbury III, Browns Mills, 


New Jersey; Dr. R. B. Dominick, McClellanville, South Carolina; Joseph 
P. Muller, Lebanon, New Jersey; Eric L. Quinter, Auburn, Pennsylvania; 
J. B. Sullivan, Beaufort, North Carolina. Ric Peigler of Clemson Uni- 
versity and Dr. F. Rindge of the American Museum of Natural History 
were helpful in arranging for me to examine those collections. Special 
thanks are due to Annie Carter of Batsto, New Jersey and to J. W. 
Cadbury III for their generous help in operating my traps, without which 
some of the records would not have been obtained. Dr. T. D. Sargent, 
University of Massachusetts generously supplied females from which ova 
were obtained and allowed me to collect extensively at his lights. An 
early draft of this report was submitted for credit in the Biology Depart- 
ment of St. Joseph’s College, Philadelphia in May 1972 (Dr. R. W. 
Fredrickson, advisor). 


AnonyMous. 1923. Records of Lepidoptera not in the New Jersey Report of 
1909. Bull. Brooklyn Entomol. Soc. 18: 136-137. 

CANADA DEPARTMENT OF Forestry. 1962. Forest Lepidoptera of Canada, Vol. 2. 
Forest Entomology and Pathology Branch Bull. 128. 

Daruincron, E. P. Notes on blueberry Lepidoptera in New Jersey. Trans. Amer. 
Entomol. Soc. 78: 33-57. 

FERNALD, M. L. 1950. Gray’s Manual of Botany. 8th ed. American Book Co. 

Forses, W. T. M. 1954. Lepidoptera of New York and Neighboring States. Part 
III, Noctuidae. New York Agr. Exp. Sta., Memoir 329, 433 p. 

FRANCLEMONT, J. G. 1942. Notes on some Cuculliinae, Phalaenidae (Lepidoptera ) 
II. On the identity of Lithophane ferrealis Grote and Xylina inominata Smith, 
with some descriptions of new forms of the genus Lithophane. Hiibner. Entomol. 
News 53: 30-35, 63-66. 

1943. On the identity of Glaea pastillicans Morrison and the species of 

the genus Chaetaglaea new genus. Entomol. News 54: 94-97. 

1968. A new species of Metaxaglaea (Lepidoptera, Noctuidae, Cucul- 

liinae). Entomol. News 79: 57-63. 

1969. Two new species of Lithophane from California ( Noctuidae, Cucul- 
limae). J. Lepid. Soc. 23: 10-14. 

Hampson, Sir G. 1906. Catalogue of Lepidoptera Phalaenae in the British Mu- 
seum. 6: 461. 

HoLianp, W. J. 1903. The Moth Book. Doubleday, New York. 479 p. 

Kimpati, C. P. 1965. The Lepidoptera of Florida. Div. Plant Industry State of 
Fla. Dept. Agri. Gainesville, Fla. 363 p. 

McCormick, J. 1970. The Pine Barrens: a preliminary ecological inventory. New 
Jersey St. Mus. Res. Rep. 2. 103 p. 

Mutter, J. P. 1965. Supplemental list of the macrolepidoptera of New Jersey. 
J. N.Y. Entomol. Soc. 73: 63-67. 

Puatr, A. 1969. A lightweight collapsible bait trap for Lepidoptera. J. Lepid. 
Sor. 23: 97-101. 

SHAPIRO, A. M. 1965. Lepidoptera records for southeastern Pennsylvania. Entomol. 
News 26: 91-95. 

Tretz, H. M. 1952. The Lepidoptera of Pennsylvania, a manual. Penn. St. Col., 
Sch. Agri., Agr. Exp. Sta. State College, Penn. 180 p. 



125 Cedar Lane, San Jose, California 95127 

Increased collecting in southern Texas in recent years has resulted in 
a number of additions to our butterfly fauna. In addition, certain species 
formerly thought to be stragglers are now known to be of regular though 
perhaps infrequent or local occurrence. Others seem to be more or less 
cyclical, present in numbers in some years, scarce or absent in others. 
The activities of many workers are adding steadily to our formerly 
meager store of information about Texas butterflies. 

I have reported on some species elsewhere (Tilden 1964; 1965a, b; 
1971). Some of these notes were recorded on trips with my good friends 
Roy and Connie Kendall, to whom I am indebted for many favors. 
Collecting in the Santa Ana Wildlife Refuge (here shortened to Santa 
Ana WLR) was under permit from the U.S. Fish and Wildlife Service. 
All specimens leg. J. W. Tilden unless otherwise stated. 


Nyctelius nyctelius (Latreille). CAMERON CouUNTy: Brownsville, 3 ¢ 6, 20 October 
1972; 3 29, 29 October 1972; 1 ¢, 1 November 1972. Males worn. A powerful 
flier; perches on tips of shrubs in the open. 

Panoquina sylvicola (Herrich-Schaffer). HmALGO couNTy: Santa Ana WLR, 1 4, 
5 November 1972; 1 4, 7 November 1972; 1 @, 9 November 1972. CAMERON 
couNTy: Brownsville, 1 9, 1 November 1972. 

Panoquina hecebolus (Scudder). CAMERON COUNTY: Route 4, 6 miles west of Boca 
Chica, 1 2, 12 October 1971. 

Panoquina evansi (Freeman). CAMERON COUNTY: Brownsville, 1 9, 20 October 
1972. HIDALGO couNTy: Santa Ana WLR, 1 6, 9 November 1972. The three I 
have taken have all been at flowers of Eupatorium odoratum, in partial shade. 

Calpodes ethlius (Stoll). Usually frequents yards, feeding on Canna, but is found 
far afield occasionally. HIDALGO couUNTy: Santa Ana WLR, 2 ¢ 6, 15-17 November 
1970, in yard, at Papaya flowers; 1 ?, 6 July 1972, on forest trail near Rio Grande. 
SAN PATRICIO COUNTY: Rob & Bessie Welder Wildlife Foundation Refuge, 1 ¢, 
on Canna in yard. 

Yoretta carus (Edwards). JEFF DAVIS COUNTY: Wild Rose Pass, 1 2, 21 June 1963; 
12 miles west of Alpine, 2 ¢ 6,1 9, 24 June 1963. 

Ancyloxypha arene (Edwards). MAVERICK COUNTY: Quemado, 7 ¢ 46, 3 2 Q, all 
slightly worn. Associated with a wet grassy roadside hollow. Kendall (1966b) has 
shown that the related A. numitor feeds on a grass (Zizaniopsis milacea) which 
grows only in wet places. The habitat of A. arene suggests that it may have 
similar habits. 

Cymaenes odilia trebius (Mabille). HmALGO couNTy: Fairly common in Santa Ana 
WLR, October-November 1970-72, on shady trails and at flowers of Plumbago 
scandens L. 

VOLUME 28, NUMBER 1 253} 

Vidius perigenes (Godman). CAMERON COUNTY: 5 miles west of Boca Chica, 2 ¢ 2, 
1 2, 30 October 1970; Brownsville, 1 @, 19 October 1972. In nature, closely 
associated with a coarse bunch grass. Has been reared by Kendall (1966a) on 
the introduced St. Augustine Grass (Stenotaphrum secundatum ). 

Monca telata tyrtaeus (Plotz). HIDALGO CouNTy: Santa Ana WLR, common October— 
November 1970, less so in 1972, on trails and in forest openings, usually in deep 

Synapte malitiosa pecta Evans. HIDALGO COUNTY: Santa Ana WLR, common shady 
trails and openings, October-November 1970, much less so in 1972. Sits quietly 
on ground for long periods of time if not disturbed. 

Pholisora alpheus (Edwards). CAMERON COUNTY: 4-6 miles west of Boca Chica 
on Route 4, 4 ¢ 46,1 2, 20 October 1963; 1 ¢, 1 9, 12 November 1963; 6 ¢ 6, 
2 22, 20 October 1970; 2 3 6, 1 2, 30 October 1970. The 1963 records seem 
to be the first for south coastal Texas. In 1972 the area was found to have been 
bulldozed and the colony destroyed. 

Gesta gesta invisus (Butler & Druce). MAVERICK COUNTY: Quemado, 1 ¢, 8 October 
1963, new county record. This species is more common on parts of the eastern 
coastal plain of Texas. 

Chiomara asychis georgina (Reakirt). SAN PATRICIO COUNTY: Rob and Bessie 
Welder Wildlife Foundation Refuge, 1 ¢, 13 October 1963, new Refuge and 
county record. Fairly common in lower Rio Grande Valley, less common westerly 
but extends to Arizona. 

Timochares ruptifasciatus (Plotz). CAMERON COUNTY: Brownsville, 1 ¢, 19 October 
1972, on lower flowers of Verbesina. 

Xenophanes trixus (Stoll). CAMERON coUNTy: Brownsville, a fair series, 20-31 
October 1972. Very inconspicuous, sitting with wings spread, on lower leaves of 
Verbesina and Eupatorium. 

Pellicia angra Evans. HIDALGO COUNTY: Santa Ana WLR, 1 4, 6 July 1972; 4 6 4, 
6 22, 16 October-18 November 1972. May be confused with the much more 
common Achlyodes thraso tamenund. 

Celaenorrhinus stallingsi Freeman. HIDALGO COUNTY: Santa Ana WLR, 1 4, 10 
November 1972, taken among undergrowth in deep shade. 

Cabares potillo (Lucas). CAMERON couNTy: Brownsville, 1 2, 30 October 1963; 
Santa Maria, 1 ¢, 18 October 1972. umaLco county: Santa Ana WLR, 1 9, 
21 October 1970; 1 9, 22 October 1970; 1 ¢, 27 October 1970; 1 ¢, 29 October 
1970; 1 9, 4 November 1970; 1 @, 13 November 1970. Occasional but widely 
scattered; visits flowers of Verbesina, Eupatorium, and Plumbago. 

Astraptes fulgerator azul (Reakirt). umDALGO couNTy: Santa Ana WLR, 1 ¢, 5 
November 1970; 1 6, 16 October 1972; 1 9, 7 November 1972. Frequents shady, 
overgrown places. 

Astraptes anaphus annetta Evans. CAMERON CouUNTY: Brownsville, 1 ¢, 18 October 
1972, at flowers of Eupatorium odoratum L. 

Chioides zilpa (Butler). HIDALGO CouNTy: Sullivan City, 1 9, badly worn, at flowers 
of Cordia boissieri DC. ( Anacahuite ). 

Phocides pygmalion lilea (Reakirt). CAMERON couNTYy: Brownsville, 4 ¢ ¢, 20 
October 1972; 1 6, 1 2, 31 October 1972; others seen, too worn to take. HIDALGO 
COUNTY: Santa Ana WLR, 1 ¢, 17 October 1970. 


Papilio anchisiades idaeus Fabricius. HIDALGO COUNTY: Santa Ana WLR, 2 é 2, 
19 October 1970, several worn individuals seen on other days. CAMERON COUNTY: 
Brownsville, 1 ¢, worn, 19 October 1972. 

Papilio astyalus pallas Gray. HIDALGO CoUNTy: Santa Ana WLR, 3 6 6, 17 October 
1972; 1 $6, 5 November 1972; 1 @ seen close-up, 7 July 1972. CAMERON COUNTY: 
Brownsville, several ¢ ¢, circa 9 October 1972, leg. Perry Glick, in his yard. 



Eurema daira lydia (Felder & Felder). HipALGo couNTy: Santa Ana WLR, 1 4, 
16 October 1972. 

Eurema boisduvaliana (Felder & Felder). HIDALGO CouNTYy: Santa Ana WLR, 1 64, 
17 October 1970. 

Eurema nise nelphe (R. Felder). Fairly common in the lower Rio Grande Valley along 
forest trails. Unlike Eurema lisa, seldom found in the open. 


Lasaia sula peninsularis Clench. Common in the lower Rio Grande Valley in October. 
Three specimens, 2 ¢ 6, 1 2, 20-22 October 1963, are aberrant, having a complete 
row of marginal light spots on both upper and lower surfaces. 

Apodemia mormo mejicanus (Behr). HUDSPETH COUNTY: near Sierra Blanca, 1 4, 
2 2 9, 20 June 1963. PREsIDIO couNTy: Shafter, 2 2 9, 7 October 1963. 

Apodemia palmerii (Edwards). CULBERSON COUNTY: Van Horn, 1 @, 23 July 1967. 
JEFF DAVIS COUNTY: 12 miles west of Alpine, 1 9, 24 June 1963. PREsIDIO COUNTY: 
Shafter, 1 ¢, 7 October 1963. 

Apodemia walkeri Godman & Salvin. CAMERON COUNTY: Brownsville, short series, 
both sexes, at flowers of Serjania brachycarpa Gray, 17-30 October 1963; South- 
most, 1 9, worn, 29 October 1963. 

Apodemia multiplaga Schaus. CAMERON COUNTY: Brownsville, 1 ¢, at flowers of 
Serjania brachycarpa Gray, 30 October 1963; 1 @, at flowers of Verbesina, 19 
October 1972. 

Calephelis rawsoni McAlpine. BEXAR CouNTY: FM 1604 at Babcock, 16 miles south- 
west of San Antonio, 1 ¢, 1 9, 11 October 1963. comax county: New Braunfels, 
4 2 2, 27 October 1972. BREWSTER COUNTY: Boquillas Canyon, Big Bend National 
Park, 1 ¢,1 2, 23 June 1963. 


Tmolus azia (Hewitson). CAMERON couNTy: 1 9, 29 October 1972, at flowers of 
Serjania brachycarpa Gray. 

Callophrys xami (Reakirt). CAMERON COUNTY: 5 miles west of Boca Chica on 
Route 4, fairly common in October 1970. This area has been bulldozed and the 
colony destroyed or greatly reduced. HmALGO coUNTY: Santa Ana WLR, 1 9°, 
on a woodland trail! 

Callophrys goodsoni (Clench). HIDALGO couNTy: Santa Ana WLR, common in 
October 1970. Very scarce in October 1972, perhaps due to dry conditions; 
a single ¢,5 November 1972. 

Strymon yojoa (Reakirt). CAMERON COUNTY: Brownsville, 1 ¢, slightly worn, at 
flowers of Eupatorium odoratum L. 

Strymon albata sedacia (Hewitson). HIDALGO COUNTY: Santa Ana WLR, 2 66,7 
November 1972, sitting on shrubbery, making short flights. See Kendall (1972). 

Strymon alea (Hewitson). COMAL CouNTY: New Braunfels, 2 ¢ ¢, 27 October 1972. 
HIDALGO COUNTY: Santa Ana WLR, 1 2, 19 October 1972. 


Apatura laure (Drury). HIDALGO couNTy: Santa Ana WLR, 1 ¢, 16 October 1972, 
sitting on Celtis pallida Torr. See Rickard (1969). 

Biblis hyperia aganisa Boisduval. BEXAR COUNTY: San Antonio, | ¢, 7 October 1968, 
leg. William Tyson. CAMERON couUNTy: 3 miles east of Brownsville, 1 ¢, 30 
October 1970. 

Dynamine dyonis Geyer. CAMERON COUNTY: Brownsville, 1 9, 13 October 1963. 

Myscelia ethusa Boisduval. HIDALGO couNTy: Santa Ana WLR, fairly common along 

VoLUME 28, NUMBER 1 25 

shaded trails under forest canopy; sits head down on tree trunks. CAMERON 
couNTy: Brownsville, 1 ¢, 2 November 1972. 

Marpesia petreus (Cramer). CAMERON COUNTY: Brownsville, 1 @, 1 November 
1972, at flowers of Eupatorium odoratum L..; two others seen same day. Broken 
weather, with short showers. 

Limenitis archippus watsoni (dos Passos). LIVE OAK COUNTY: North end of Lake 
Corpus Christi, 2 6 6, 1 9, 12 October 1963; 3 ¢ 6, 2 99, 7 November 1963. 
Very similar to specimens from Louisiana. 

Metamorpha stelenes biplagiata (Fruhstorfer ). HIDALGO COUNTY: Occasional in Santa 
Ana WLR, usually badly worn; 2 nearly perfect ¢ ¢, 27 October 1970. CAMERON 
COUNTY: Santa Maria, 1 worn @, 18 October 1972; Brownsville, seen on three 
occasions in October 1972, all too worn to collect. 

Anartia fatima (Fabricius). CAMERON COUNTY: Santa Maria, 2 646, 1 9, 18 
October 1972; taken also by the Kendalls at the same time and place. HIDALGO 
coUNTY: Santa Ana WLR, 3 ¢ 6, 4 July 1972; Bentsen-Rio Grande Valley State 
Barks MOS July 1972. 

Junonia evarete (Cramer). HIDALGO cCouNTy: Santa Ana WLR, 1 64, 15 October 
1970, leg. Wayne Shifflett. CAMERON couUNTy: Brownsville, 1 ¢, 21 October 
1963. These may represent subspecies zonalis Felder & Felder. I have seen other 
specimens. Sympatric with Junonia coenia in the lower Rio Grande Valley. 

Junonia nigrosuffusa Barnes & McDunnough. NUECES couNTy: Mustang Island, 1 
6, 15 October 1963; others seen. I have seen a number of specimens from coastal 
Texas, taken by others. 


KenpaLL, R.O. 1966a. Larval foodplants for five Texas Hesperiidae. J. Lepid. Soc. 
20: 35-42. 

. 1966b. Larval foodplants and distribution for three Texas Hesperiidae. 

J. Lepid. Soc. 20: 229-232. 

1972. Three butterfly species (Lycaenidae, Nymphalidae, and Heliconi- 
idae) new to Texas and the United States. J. Lepid. Soc. 26: 49-56. 

Rickarp, M.A. 1969. In, News Lepid. Soc. No. 3, Annual Summary, p. 12, line 30. 

TILDEN, J. W. 1964. Two species of Hesperiidae previously unrecorded from the 
United States. J. Lepid. Soc. 18: 214-216. 

1965a. Urbanus procne and Urbanus simplicius (Hesperiidae). J. Lepid. 

Soc. 19: 53-55. 

1965b. The genus Panoquina occurring in Texas. J. Res. Lepid. 4: 37-40. 

. 1971. Aguna claxon (Hesperiidae) new to the United States. J. Lepid. 

Soc. 25: 293. 



Joun R. G. TuRNER? 
Department of Biology, University of York, England 

Heliconius are among the most attractive of South American butterflies; 
they have beautiful bright color patterns, and their elongated wings 
enable them to perform quite remarkable tricks in flight, such as hover- 
ing, vertical climbs, and even flying backwards over short distances. 
Investigations on various aspects of their biology, reported in a wealth 
of papers, mainly in Zoologica and the Journal of Insect Physiology, have 
also made these animals into important research tools in such diverse 
fields as evolutionary genetics, ecology, behavior, and physiology. This 
is a brief account of the methods required to culture these organisms out- 
side their normal tropical environment. 

The methods described are those developed for genetical work on 
three species (H. melpomene, H. erato and H. charitonia) in England. 
Other species sometimes require rather more space, and, of course non- 
genetical work requires much less separation of females and therefore 
lends itself much more to mass culture. The results of the genetical 
experiments will be reported elsewhere (Sheppard and Turner, in prep.; 
Turner, 1973). 

Techniques for culturing Heliconius in the tropics were developed 
under the guidance of William Beebe and Jocelyn Crane in Trinidad and 
have been described fully elsewhere (Crane & Fleming, 1953; Turner & 
Crane, 1962). 

Culturing Adult Butterflies 

Heliconius need to be kept at a temperature between 70°F and 105°F; 
below about 68°F they tend to become inactive (a slightly higher 
temperature for equatorial races, a slightly lower one for temperate 
races), but the night temperature when the butterflies are roosting can 
be brought as low as 60°F without obvious ill effects on the stock. Tem- 
peratures over 105°F become very dangerous after some time, particularly 
for butterflies which are already in a physiologically weakened con- 
dition, and in a changeable climate it is advisable to have thermostatically 
controlled windows as well as thermostatically controlled heating of 
the greenhouse. 

1 Dedicated to the memory of Roni Grainger. 
2Present address: Department of Ecology & Evolution, Division of Biological Sciences, State 
University of New York at Stony Brook, Stony Brook, New York 11790. 

VoLuME 28, NUMBER 1 2h 

A greenhouse provides the simplest way of producing an agreeable 
environment for the butterflies, as it can be relied on to produce diurnal 
fluctuations in conditions without resort to the elaborate programming 
machinery required when using an enclosed artificial environment cham- 
ber. However, during the winter the surface of the glass becomes 
extremely cold and the butterflies must be kept off it, either by double 
glazing or by enclosing the butterflies in some kind of mesh cage within 
the greenhouse. The minimum size of cage in which the butterflies will 
breed normally and live out a reasonable span has not been accurately 
determined. Breeding experiments in York, England were conducted in 
walk-in cages about 9 X 8 X 7 ft. and most forms did pretty well in 
these, although there was some obvious variation: the South Brazilian 
race of H. erato bred extremely well; the Amazonian race of H. melpo- 
mene bred well, although some individuals behaved as if they were a 
little unhappy in the confined space; hybrids between the South Brazilian 
and Amazonian races of H. melpomene spent excessive amounts of time 
on the roof of the cage and showed signs that a larger chamber might 
have served them better. 

The London Zoo succeeded in breeding the Amazonian race of H. 
melpomene in a cage about 3 X 3 X 4 ft., but this is probably only to 
be recommended when space is very short. With care it is also possible to 
breed from female butterflies kept in the sunny bay window of a sitting 
room with good background central heating, but only about one in five 
females takes to this life sufficiently to live more than a week or so, 
and to lay eggs. 

Temperate variations in day length do not seem to upset the behavior 
of the butterflies unduly, nor does cloudy weather, with the exception 
of very thick cloud during the winter. On the whole roosting takes 
place at roughly the normal time of tropical sundown, but for cloudy 
weather it may be worth providing some artificial light in the form of 
strip lighting suspended over the cages. It is a mistake to place the 
lighting inside the cage as the butterflies tend to damage themselves 
by flying against the elaborate fittings. 

Cages are best constructed from a bolted frame of pre-drilled angle 
iron, which is commercially available, covered with mosquito net or 
“Lumite” saran screening fixed on with a rubber adhesive. Sliding doors 
are quite easy to construct with such material, and should be no more 
than waist high, to reduce the possibility of butterflies escaping when 
the doors are opened; an extra curtain of netting hanging loose across the 
inside of the door is an added insurance. Butter muslin (cheesecloth) 
is not recommended for cages as it is difficult to see what is going on 


Figs. 1-2. The Heliconius breeding-system used at the University of York: (1) the 
greenhouse. Passiflora plants to the left and upper right, a pair of cages to the right 
of the door. (2) a pair of cages in use. To the left inside the cage—Passiflora serrato- 
digitata and Abutilon sp.; in the background—Passiflora caerulea. The slits on the 

left and in the wall separating the two cages are for introducing plants on long canes. 
(Photographs by Richard Hunter. ) 

VOLUME 28, NuMBER 1 29 

inside the cage, thus cutting down attention to emergencies, and poly- 
thene is likely to produce unfortunately stagnant conditions in the air in 
the cage. An alternative of course is to buy commercially manufactured 
cages. (For the use of a plastic netting tent, see the article by J. Brewer, 
News of the Lepidopterists’ Society, 1972, number 6.) 

Heating elements should be kept out of cages, as butterflies, par- 
ticularly when sick, can destroy themselves by landing on them. 

If humidifiers are not available, then high humidity may be main- 
tained in the cages by frequent spraying of the floor with a hose. In 
a greenhouse with a concrete floor it pays to cover this with heavy duty 
polythene sheeting, as this collects puddles. To act as a more constant 
supply of humidity, particularly during hot weather, it helps to have a 
bed of saturated peat about six inches deep occupying about half the 
floor of each cage. 

Heliconius butterflies take pollen in addition to nectar (Gilbert, 
1972). Food sources are therefore a flowering plant which produces a 
plentiful supply of pollen (I have found Abutilon excellent for this 
purpose and readily obtained), and failing nectiforous flowers, a supply 
of honey. I have found it best to supply honey neat on the petals of a 
plastic flower, and also diluted in water in a dispenser of the type used 
for giving drinks to caged birds. It is also good to have a supply of pure 
water dispensed from a wet sponge, in addition to the supply of puddles 
on the floor. Because of the preferences for red or orange flowers shown 
by most Heliconius (Crane, 1955) it is good if sponges, plastic flowers and 
bird feeders are of one of these colors. In winter, honey can be changed 
once every two or three days, but in hot weather daily changes are 
needed, particularly of the honey-water mixture, to reduce the concen- 
tration of alcohol. 

The cage should be supplied with additional plants to provide perch- 
ing surfaces for the butterflies, and also shade; moving out of hot dry 
areas seems to be an important factor in the survival of the butterflies 
in the greenhouse during hot weather. Grevillea robusta is excellent for 
this purpose, and provides a photogenic background. 

Culturing the Early Stages 

Heliconius larvae feed on quite a wide variety of the five to six hun- 
dred species of Passiflora (see e.g. Alexander, 1961; Brown & Mielke, 
1972). The three species used in the present experiment all laid and fed 
readily on Passiflora caerulea, which is a very vigorous grower and can 
be obtained easily from nurserymen in England (not in the USA), as 
it is a popular ornamental. Another good hardy species with large leaves, 


although a little slower growing, is the horticultural hybrid P. allardi 
(again unfortunately very rarely cultivated in America), which is eaten 
readily by melpomene and erato at least. In addition any of the tropical 
species which are natural foodplants (for example P. laurifolia and P. 
serrato-digitata for melpomene) may be used, but tend to be much 
slower growing and therefore harder to replenish. P. biflora makes a 
good foodplant for erato, but suffers badly from exposure to sun and 
low humidity. 

Females lay regularly on the growing shoots of Passiflora vines placed 
in the cages. The simplest technique is to place a healthy young plant, 
potted and on a six foot cane, in the peat bed along with the other plants. 
The larvae can be left to feed on this plant, and require little attention, 
as cultures of these species outside the tropics seem to be relatively free 
of epidemic diseases. However, the larger larvae are prone to eat the 
young growing shoots of the plant; as these are the only sites used by 
the females for laying, they rapidly slow down the rate of egg production, 
as well as destroying any eggs and young larvae that are on the shoots 
when they are consumed. It therefore pays to move half-grown larvae by 
hand to the lower and older parts of the plant. With judicious transfers 
of larvae, three healthy caerulea can keep pace with the offspring of a 
normally fertile Heliconius, and thus provide one with continuous culture. 
For non-genetical work where the offspring of several females are mixed 
it is clearly necessary to provide more plants. 

With an adequate food supply the larvae of the different species do 
not seem to compete excessively, and all can be cultured in the same cage. 
The larvae of H. erato and H. charitonia pupate on or near the plant on 
which they have fed and can safely be left to do so. H. melpomene 
larvae tend to wander between six and twelve feet before pupation 
(the warning sign of this is that they turn bright purple) and may thus 
get into the wrong cage, producing contamination of another brood. 
This is avoided by placing final instar larvae in standard cylindrical 
breeding cages (obtainable from English suppliers), to feed on cut vine 
stalks. The slow-growing but tough-leaved P. laurifolia is ideal for this 
purpose. The larvae will then pupate either on the gauze lid of the 
cylinder or on the cut stalks. Once all the larvae have pupated the stalks 
are placed vertically in the peat bed, and the butterflies allowed to eclose 
freely in the cage. 


Figs. 3-6. The Heliconius breeding-system used at the University of York: (3) 
To show the sliding door of the cage. On the left, Passiflora serrato-digitata and P. 
auriculata. (4) Inside the corner of a cage, showing the peat bed, polythene-covered 

VoLUME 28, NUMBER 1 31 

floor, butterfly-feeders, Abutilon (left), Grevillea (center), P. caerulea (foreground, 
mostly defoliated). (5) Transferring Heliconius melpomene larvae to a cylinder 
for pupation. Buckram cylinders behind. (6) An interracial hybrid of Heliconius 
melpomene feeding from a honey-water dispenser. (Photographs by Richard Hunter. ) 


Pupae which have become detached from their silk pad for any reason 
may be placed to eclose in the bottom of one of the larval breeding 
cylinders in which the plastic walls have been replaced by a cylinder 
of stout buckram, but this method results in the crippling of about one 
butterfly in five, when it fails to climb the buckram to blow out its wings. 
It is better, but more time consuming, to stick the cremaster of the pupa 
onto a woody branch, using a little clear rubber adhesive. This method 
has a high success-rate. 

Eclosed offspring are collected and frozen once every one or two days, 
or transferred to other cages for breeding, or of course may be left in 
the cage for a continuing culture. The delay of a few days before males 
become fertile after eclosion gives one a little latitude in collecting them. 

Butterflies are best transferred from cage to cage not in the hand, 
which may injure them, but in the small suspended gauze cages which 
breeders use for mating large silk moths. 

When a female dies or is killed and her cage is required for another 
brood, the existing Passiflora plants can be covered completely with 
black organdie (organza) sleeves to separate the old brood from the new. 


Princes or professional researchers might seem to be the only people 
with the resources to grow these butterflies. Certainly genetical work 
requires a large amount of space because it is necessary to separate each 
female in a six foot or larger cage; in addition a considerable amount of 
greenhouse space is taken up with the stocks of Passiflora vines. But there 
is no reason why an amateur with a reasonably well appointed green- 
house, particularly in the warmer parts of the temperate zone, should 
not be able to cultivate these insects for fun. This they certainly provide. 


I am grateful to Miss Jocelyn Crane-Griffin, who instructed me in 
Heliconius-technique in Trinidad, and to Professor Philip M. Sheppard 
FRS, who originated several of the greenhouse techniques described here. 

The detailed design of the cages, and their construction, was the work 
of the late Miss Veronica A. Grainger. 

The work would not have gone forward without the expert attention 
given to the Passiflora plants by Mr. Colin Abbot and Mr. John Arber. 

The final preparation of this paper was supported by a Biomedical Sci- 
ences Support Grant (HEW Grant #5805 RR 07067) awarded to the State 
University of New York at Stony Brook and by NSF grant #B039300. 



VoLUME 28, NuMBER 1 pre) 


ALEXANDER, A. J. 1961. A study of the biology and behavior of butterflies of the 
subfamily Heliconiinae in Trinidad, West Indies, Part I. Some aspects of larval 
behavior. Zoologica (New York) 46: 1-24. 

Brown, K. S. & O. H. H. Mietxe. 1972. The heliconians of Brazil (Lepidoptera; 
Nymphalidae). Part II. Introduction and general comments with a supple- 
mentary revision of the tribe. Zoologica (New York) 57: 1-40. 

Crane, J. 1955. Imaginal behavior of a Trinidad butterfly, Heliconius erato hydara 
Hewitson, with special reference to the social use of color. Zoologica (New 
York) 40: 167-196. 

& H. FLreminc. 1953. Construction and operation of butterfly insectaries 
in the tropics. Zoologica (New York) 38: 161-172. 

GitperT, L. E. 1972. Pollen feeding and reproductive biology of Heliconius 
butterflies. Proc. Nat. Acad. Sci. USA 69: 1403-1407. 

TurNER, J. R. G. 1973. Passion flower butterflies. Animals 15: 15-21. 

& J. Crane. 1962. The genetics of some polymorphic forms of the butter- 

flies Heliconius melpomene Linnaeus and H. erato Linnaeus. I. Major genes. 

Zoologica (New York) 47: 141-152. 


126 Wells Rd., Hanahan, South Carolina 29405 

From my field observations during the past four years, along with data 
gathered from other sources, I have come to certain conclusions regarding 
the breeding habitats, flight habits and foodplants of Satyrium kingi 
(Klots & Clench). 

I first collected Satyrium kingi in Escambia County, Florida, near 
Cantonment. It was a single worn female, taken 2 August 1969. I was 
not sure of its identification until I compared it with material I collected 
in South Carolina in 1970. 

In addition to the Florida spot, I have taken kingi at four localities 
in South Carolina: (1) at Givhans Ferry State Park in Dorchester 
County; (2) on the south side of Highway 642 where Dorchester and 
Charleston Counties meet, about 200 yards inside Dorchester County; 
(3) in Berkeley County at the Naval Weapons Station, near the golf 
course; and (4) in Charleston County just outside the south gate of the 
Air Force Base along Dorchester Road. 


These five localities can be divided into two types. One type, including 
the Florida locality and the two in Dorchester County, South Carolina, 
will be referred to as group A. The other two localities are called 
group B. 

The group A localities are wooded areas, with few or no flowers at 
the time kingi was taken. Although I have not personally observed 
ovipositing in any of the 40 or so females I have collected, nor have any 
oviposited after capture, still it is my belief that sweet gum ( Liquidambar 
styraciflua) will prove to be a primary foodplant of S. kingi. Ninety 
percent of the females which were not taken at flowers were collected 
from the leaves of sweet gum saplings. The other ten percent were on the 
leaves of various other plants near sweet gum. 

In all the group A localities kingi was found around the edges of old 
forests where sweet gum saplings grew. I believe that much the same 
situation may exist with S. kingi as with Papilio aristodemus ponceanus 
(Schaus), where the species depends on second growth forests around 
older forests as its habitat (Rutkowski, 1971). Man’s efforts may hurt 
ponceanus by overprotecting hammocks from natural disaster such as 
fire or storm, or by clearing hammocks away for construction, but his 
works may actually help kingi by cutting roads, making fire lines and 
power line cuts through old or virgin forests, thus providing areas for 
sweet gum saplings to grow. 

I first found kingi in South Carolina at the Highway 642 locality in 
1970. That season I collected some 15 females but only one male (on 
Holly (Ilex) ) at that locality. During the 1971 season I again collected 
female kingi at this spot and also took a very few specimens from the 
Givhans Ferry State Park, type A locality, all females. In 1972 the High- 
way 642 locality again yielded several females but no males. By this 
time I had noticed the marked affinity the females had for sweet gum. 
So when I visited the Givhans Ferry State Park spot in 1972 I moved my 
collecting efforts 20 yards from where I had collected the year before 
and found female kingi in good numbers in a stand of sweet gum saplings. 
However, I had still not found any more males in the type A areas in 
three years! , 

In June 1972, at the Givhans Ferry State Park locality, I noticed a 
small butterfly come darting from high in the top if a mature sweet gum 
down to the young saplings where I was collecting female kingi, then 
return to the taller trees. A little later either the same specimen or 
another one did the same thing. However, before it could fly up again 
I netted it and found it was a male kingi. A little later I noticed two 
males dart down in this fashion from their lofty perches then fly up again 

VoLUME 28, NuMBER 1 35 

accompanied by females (this occurred at about 1630 EDT). I never 
saw a pair in copula, but did observe their courtship flights. Female 
kingi fly fairly low, from 4 to 10 ft. above the ground, around sweet gum 
saplings in the type A areas. Males stay high in older trees darting down 
to the saplings to look for females. After finding a female, the male 
accompanies her in a flight nearly straight up into the older trees. 

The group B localities are open areas with tall flowering hedges. 
These flowering hedges were three to four hundred yards from any 
forests, but a limited number of kingi were taken there of both sexes. 
Obviously, they were there simply as flower visitors, and not because of 
any suitable breeding habitat. In the B areas, male and female kingi 
fly in more equal numbers, with males being slightly more numerous. 
Adults were never observed investigating each other or engaging in 
courtship flights. In 1971 and 72 twelve specimens were taken from the 
B areas, eight of which were males. 

A significant distinction between the group A and B areas is that 
although both males and females were taken at group B spots, the vast 
majority of specimens taken from group A areas were females, with 
only two males collected there. The great contrast between the A and 
B areas leaves little doubt that kingi was in the B areas only to visit 
flowers. The absence of flowers in the A areas along with the large num- 
ber of specimens, mostly females, found there year after year is strong 
evidence that kingi breeds in the A areas. 

Even in the A areas, which seem to be the preferred natural breeding 
habitat, kingi is local in occurrence. It is found most commonly only 
where sweet gum saplings grow, and rests on the leaves of this plant. 
S. kingi shows such a marked affinity for sweet gum that this tree is 
presumably a foodplant. I have never found kingi in much searching in 
any other type of habitat, except when visiting flowers. 

The place to look for kingi is around the edges of old, well established 
forests. These may be mixed hardwood and pine forests or hardwood 
alone. The butterfly may be found whether these forests are in low 
swampy areas or rather dry areas going into open pine flats and forest. 

Males stay high in older trees, not always sweet gum, except when 
visiting flowers, at which time they are easily caught. Even when visiting 
flowers kingi males like the higher blossoms. Female kingi do not seem to 
visit flowers as much as males. Females should be looked for on sweet 
gum samplings usually only 5 to 6 ft. from the ground. 

All five of the areas where I have collected kingi represent coastal popu- 
lations. Satyrium kingi was described in 1952 from the coastal population 
at Savannah, Georgia, by Klots and Clench. In the original description, 


under the heading “Ecological Data,” there are several items of interest. 
First, the Dorchester County locality in South Carolina can be described 
in almost the exact words used to describe the type locality (Klots & 
Clench, 1952, p. 15). Second, sweet gum (Liquidambar styraciflua) is 
noted as one of the plants found at the type locality. Third, as the 
collecting in the type locality was done while specimens were visiting 
flowers, more males should have been taken than females, and indeed 
the type series consisted of 5 males and 2 females, taken over a period 
of three years. It is also noted in the original description that specimens 
preferred the higher flowers they were visiting. 

When describing kingi, Klots and Clench mentioned 5 specimens not 
included in the type series. These specimens were excluded from the 
type series “because of the danger of future subspecies confusion.” Here 
in the original description a very important distinction is made between 
typical lowland (coastal) kingi and the inland or highland populations. 
(Klots & Clench, 1952, p. 8.) Klots and Clench saw the likelihood that 
northern inland populations of kingi might represent subspecifically 
distinct populations. 

Mr. Lucien Harris Jr. in his recent book, “Butterflies of Georgia,” stated 
that several years after kingi was described he too gave some thought 
to naming a subspecies from the inland areas of Georgia, but he decided 
to leave this to the “experts in this field.” 

If there is a subspecies involved in these highland populations the 
choice of foodplant is important. Because of this question, the relation of 
S. kingi, in my experience, to old forests and sweet gum, and its flower 
visiting and courtship flight habits, seem very relevant. 

The northern (inland) population of kingi has been reared on Flame 
Azalea (Rhododendron calendulaceum) by Mr. John C. Symmes in the 
Atlanta, Georgia, area (Harris, 1972). Harris also notes that when 
H. L. King, for whom the species is named, collected kingi at the type 
locality he saw females ovipositing on a plant not related to Azalea, and 
that King did not find Azalea plants there. The ecological differentia 
between the lowland populations of kingi in Florida, Georgia and South 
Carolina, and those populations of inland and northern areas, along with 
the superficial differentia of those populations should be examined more 
closely by the experts, in my opinion. 

Coastal kingi shares its habitat with a rather small number of butterflies. 
These species are rather uncommon and are usually considered good 
catches: Autochton cellus (Boisduval & Le Conte), Poanes yehl 
(Skinner), Amblyscirtes aesculapius (Fabricius), Papilio palamedes 
(Drury), Satyrium calanus (Hibner), Satyrium liparops (Le Conte), 


Asterocampa alicia (Edwards) (following Reinthal, in Harris, 1972), 
Asterocampa clyton (Boisduval & Le Conte), Lethe creola (Skinner), 
Lethe portlandia (Fabricius), Lethe appalachia (Chermock), and 
Euptychia gemma ( Hiibner). 

In the highland and inland areas kingi is on the wing in July and 
August. In the coastal areas of South Carolina and Georgia it flies 
in May and June. The late date of the one specimen I collected in coastal 
Florida (Aug. 2) may very well mean that kingi is double brooded there. 


EuruicH, P. R. & A. H. Exruicu. 1961. How to Know the Butterflies. Brown, 
Dubuque, Iowa. 

GaTRELLE, R. R. 1971. Notes on the occurrence of two rare Lepidoptera in South 
Carolina. J. Lepid. Soc. 25: 143. 

Harzis, L., Jr. 1972. Butterflies of Georgia. Univ. Oklahoma Press, Norman. 

Kuots, A. B. & H. K. CLencH. 1952. A new species of Strymon (Huebner) From 
Georgia (Lepidoptera, Lycaenidae). Amer. Mus. Novit. 1600: 1-19. 

pos Passos, C. 1970. A revised catalogue with taxonomic notes on some Nearctic 
Lycaenidae. J. Lepid. Soc. 24: 26-38. 

Rutkowski, F. 1971. Observations on Papilio aristodemus ponceanus ( Papilion- 
idae). J. Lepid. Soc. 25: 126-136. 


As a result of the recent election, it is a pleasure to announce that Norman D. Riley 
was overwhelmingly approved by the membership as an honorary life member of the 
Lepidopterists Society. The newly elected officers of the Society are listed inside the 
front cover. In addition, Dr. W. Donald Duckworth was elected as the Jordan Medal 
Representative, and the proposed constitutional amendments (see Vol. 27, p. 241) 
were passed. 



1625 Roma NE, Albuquerque, New Mexico 87106 

Since 1964 I have resided in Albuquerque, New Mexico, and devoted 
much time to investigating the butterflies and moths of the surrounding 
mountains. The purpose of this article is to describe some of the obser- 
vations made during these nine seasons. 


Albuquerque itself is located a mile above sea level at the crossing of the Rio 
Grande Valley and U. S. Highway 66. The Valley is about 25 miles wide through- 
out this part of the state. About 75 miles north of Albuquerque, the New Mexican 
extensions of the high Colorado ranges begin dropping off into desert country. These 
northern mountains typically have 12,000—14,000 ft. peaks and 7000-8000 ft. valleys. 
However, the lower desert surrounding Albuquerque itself is only 4000-7000 ft. 
in elevation. 

Scattered ranges rise from the desert at numerous points around Albuquerque (Fig. 
1). These mountains are on both sides of the Rio Grande Valley, and have peaks 
of up to 11,000 ft. On the east side of Albuquerque, the Sandia Mts. begin at the 
edge of town; a bit southeast of Albuquerque, the Manzano Mts. begin. U. S. High- 
way 66 and Tijeras Canyon partially isolate the Sandias from the Manzanos. The 
bottom of the Albquerque side of the Sandias and Manzanos is about 6000 ft. 
and fairly arid: about 8 in. of rain falls each year, mostly in the form of summer 
thundershowers. The bottom of these mountains on the side opposite Albuquerque 
averages 7500 ft., and is much more moist. Rainfall at the bottom on that side 
probably approaches 20 in. annually. To the east of the Sandias and Manzanos 
the Great Plains begin, and roll across the shallow Estancia and Pecos Valleys, 
gradually sloping down to 4000 ft. at the Texas state line. Peaks in the Sandias 
and the Manzanos are around 10,000 ft. The upper portions of both these ranges 
are heavily forested, but have Canadian zone meadows at the very summits. 

To the west of the Rio Grande, the New Mexico desert rises gently up to 7000 
ft. or 8000 ft. on the Continental Divide east of the Arizona line. In this area in 
the central latitudes of New Mexico, six or seven mountain ranges project above the 
desert to exceed 9000 ft.; the four easternmost are Mt. Taylor, Ladron Peak, the 
Magdalena Mts., and the San Mateo Mts. The high desert west of Albuquerque 
is considerably drier than the Great Plains to the east. 

Mt. Taylor is a fairly recent volcano of 11,000 ft. surrounded by a vast level 
plateau, especially to the northeast, which is uniformly just over 8000 ft. Most of this 
plateau is at least lightly forested. The top of Mt. Taylor proper breaks out into a 
large pseudo-arctic meadow of several square miles extent. Ladron Peak is a small, 
rugged monument which juts to just above 9000 ft. from the very low surrounding 
desert floor of 5000 ft. Ladron Peak is completely unforested, although the upper 
gorges with a northern exposure have scattered aspen (Populus tremuloides Michx. ) 
and ponderosa (Pinus ponderosa Lawson). The Magdalena Mts. tend to resemble 
the Sandias and Manzanos, except that they run east-west instead of north-south 
and are ower Sonoran rather than Upper Sonoran at the base. Large Canadian- 

VoLUME 28, NuMBER 1 39 

zone meadows also cover the tops of the Magdalenas, which reach nearly 11,000 ft. 
The San Mateo Mts. are isolated from the Magdalenas by the very arid Mulligan 
Valley and unpaved N. M. Route 107. They are less isolated from the Gila Mts. 
of southwestern New Mexico, at least in terms of possible dispersal routes not 
interrupted by broad, dry valleys. 

Generally speaking, each range has been studied by intensive collect- 
ing for one season, preceded and followed by two seasons of occasional 

The Colorado mountain extensions into northern New Mexico—the 
Jemez and Chuska on the west of the Rio Grande and the Sangre de 
Cristo on the east—have not been included in this work. I hope to 
examine these ranges in detail in future years. Also, the high Sacramento 
Mts. and the Guadalupe Ridge in southeastern New Mexico; and the 
Gila, Black Range, Animas, Zuni, and Datil mountains in the southwest 
are areas I want to probe eventually. Finally, the eastern plains and 
the Rio Grande riverbottom remain to be examined. (Obviously our 
very few resident New Mexican collectors will require quite a number 
of seasons to come to know a state nearly the size of California with a 
very complex life-zone pattern. For this reason, I would greatly appre- 
ciate correspondence with any outsiders who have done serious collect- 
ing here. Perhaps, with the help of outsiders, our study of the 12 
previously-mentioned virgin areas can be greatly accelerated. ) 

Abundance Tabulation 

My findings concerning occurrences and abundances of butterfly 
species in the six mountain ranges surrounding Albuquerque are sum- 
marized in Table 1. In this table, the following symbols and abbreviations 
are used: 

TL—Mt. Taylor 

SD—Sandia Mts. 

MZ—Manzano Mts. 

LD—Ladron Peak 

MG—Magdalena Mts. 

SM—San Mateo Mts. 
A—abundant (over 100 per hour ) 
C—common (over 15 per year ) 
U—uncommon (2-15 per year ) 
S—single record 
V—visual record 
X—insufficient observation to distinguish between S, U or C 
Q—record not observed by author 


?—of uncertain determination or unknown collector 
MT—Mike Toliver, collector 
KR—Kilian Roever, collector 
HK—Harry Clench, collector 
JB—John Burns, collector 
OS—Oakley Shields, collector 
!—major range extension 
(M )—migratory, either solitary or in numbers 
(L)—intensely local 
(D)—species often found on desert floor, away from mountains 
and permanent water 

Species numbers and names are as given by dos Passos (1964), except 
where revised: Philotes (Beuret, 1958) and (Langston, 1969); Melitaeinae 
(dos Passos, 1969); Vanessa (Field, 1971); Erynnis (Burns, 1964); 
Cercyonis (Emmel, 1969); Theclinae (dos Passos, 1970); and Mega- 
thymidae (Freeman, 1969). 

Correlation Coefficients of Species with Ranges 

On the map (Fig. 1) and in Lines 1 of Table 2, correlation coefficients 
px are shown of the species occurrence in the six ranges studied. These 
coefficients are computed by assigning a value X;,; of 

1 to species i if it occurs in range j, 

0 to species i if it does not occur in j, 
and then calculating: 

pi Day Xu —2/N) (X= ma/N) 

(fe = 

N N 
l . 2T 12 
Rees 2 n/N)? | | Sy (Xie—me/N)? | 

i=l i=l 
Here, nj (= 3;X;;) and nx (= 3;Xi,) are the total number of species 
occurring in ranges j and k, respectively, while N is the total number of 
species found in all six ranges. If ranges j and k have exactly the same 
species, pj, would be 1. If all the species which occur in either range 
do not occur in the other, pj, would be —-1. If one were to release in a 
room two live specimens of each of 100 species, and then permit two 
collectors to catch 100 random specimens each, p;, between collectors 
j and k would be, on the average, 0. Thus, the p;, are measures of the 
faunal similarities of the different ranges. The surprising aspect con- — 


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Fig. 1. Map of study area, showing mountain areas over 8000 ft. Numerical 
values indicate correlation coefficients of species occurrence. 

VoLUME 28, NUMBER 1 49 

cerning the species distribution correlation coefficients appearing on 
the map is their very low values. Even for the Sandias and the 
Manzanos, which are only 20 air miles apart with a 7000 ft. “bridge” 
connecting them, pjx is just 0.3. This is about the correlation coefficient 
which relates the species of New York and South Carolina! 

Several alternative procedures were also used for computing species 
distribution correlation coefficients. First of all, pj, was re-evaluated 
by letting 

Xi; = 4 if species i is abundant (A) in range j, 

3 if species i is common (C) in range j, 

2 if species i is uncommon (U or X) in range j, 

1 if species iis of dubious occurrence (S, V, X, Q or ?) in range j, 
0 if species i is unreported from range j. 

Lines 2 of Table 2 give the results of this computation. Next, pj, was 
determined by the same process, but with X;; reduced to | for all species 
indicated in Table 1 as showing tendencies to migrate or to reside on the 
desert. The effect of this modification on X;; is to de-emphasize the 
occurrence of free-moving species in computing the correlation coeffi- 
cients. Lines 3 of Table 2 show the values of p;; found by this altered 
assignment of Xjj. 

Additionally, p;; was calculated with X;; = 0 for all desert or migratory 
species. Here, the contribution of free-moving species is totally eliminated 
from the faunal similarity measurement. Lines 4 of Table 2 give these pj. 

From the results shown in Lines 1-4 of Table 2, the following general- 
izations may be drawn: 

a. Lines 4 are always greater than Lines 1-3. In statistical terminology, 
this means total suppression of frequency data on free-moving species 
gives the highest species distribution correlation coefficients. In other 
words, in comparing the characterizing fauna of isolated areas, it is best 
to ignore records of species which may frequently cross the isolating 
barriers. To do otherwise, at least on a sampling period of only nine years 
injects short-term dispersal effects. As these short-term effects are 
apparently fairly random, they cause the fauna of the various areas to 
appear more distinct than they truly are. 

b. Lines 4 are always greater than Lines 3. Statistically, this means 
partial suppression of frequency data on free-moving species gives lower 
species correlations than total suppression of these data. In simpler 
language, if one wants to ignore effects of short-term dispersal in mea- 
suring faunal similarity, one should exclude totally the records of free- 
moving species. They should not be included with emphasis merely 


TaBLE 2. Correlation coefficients of butterfly species with mountain range. 


MT 214 207 .203 395 .296 
.253 383 162 376 310 

233 361 112 375 348 

356 465 233 459 448 

761 813 656 .766 .656 

SD 332 LZ 398 309 
003 307 433 340 

04 .260 A493 oT 

.660 381 586 A81 

880 798 TAT 759 

MZ .167 304 .267 
282 ATS» ~—A28 

.204 O24 A48 

393 {5)SL7/ 063 

.768 783 778 

LP 436 344 
O41 AAT 

460 368 

44 A72 

862 S11 

MG 21 




Lines 1: Xi; constrained to 1 or O for all species. 

Lines 2: Xi; variable from 4 to O for all species. 

Lines 3: Xi; constrained to 1 or O for desert-migratory species, otherwise vari- 

able from 4 to 0. 

constrained to 0 for desert-migratory species, otherwise variable 

from 4 to 0. 

Lines 5: Xi; variable from 4 to 0 for desert-migratory species, otherwise con- 
strained to 0. 

Lines 4: X; 


c. Lines 2 are usually greater than Lines 1. Mathematically, this 
means data on abundance of species in different areas will correlate more 
highly than a simple yes-no declaration as to occurrence of each species 
in each area. Ecologically, this is a rewording of the phenomenon that 
an organism which is common (successful) in one area is likely to be 
common in another area if it is found there at all. 


d. Ladron Peak fauna correlates highly with only the Magdalena 
Mountains, although the converse is not true. Thus, Ladron Peak is 
biologically a depauperate island of the Magdalenas. 

e. The Manzano Mts., which have the most “endemic” records, do not 
have noticeably low faunal correlations with the other ranges. A situa- 
tion such as this may develop when foreign insects may drift into a new 
area more easily than they can drift out, or when an area has an unusually 
diverse foodplant flora relative to other areas in the study. 

f. Early in this study, it was anticipated that more collecting in each 
range would raise the species distribution correlations. This anticipation 
seems not to have been borne out by annual re-evaluation of the 

Correlation coefficients were also computed with data suppressed on 
desert species but not on migratory species. No unexpected trends 
appeared. Reversing desert and migratory species in this procedure 
also produced no surprises. 

As a final computation, correlation coefficients were evaluated for the 
desert and migratory species only: the X,; were reduced to zero for 
all other species. These values are given in the last lines of Table 2. 
In each range except Ladron Peak, about % of the observed species are 
considered either desert or migratory. For Ladron Peak, the fraction is 
%. One can see that, as expected, free-moving species have much higher 
correlations than other species. This statement is, in fact, almost a 
tautology: Species which can readily cross barriers are more likely to 
turn up on both sides of the barriers than species which cannot. 


I have described briefly the topography of the area around Albu- 
querque, New Mexico, and presented a table which summarizes the 
occurrence and abundance of butterflies in six surrounding mountain 
ranges as observed over nine seasons. The data included in this table 
enable computation of species-distribution correlation coefficients be- 
tween ranges. These coefficients are measures of the faunal similarities 
of the butterfly populations in the six ranges. 

Appendix: Notes on Callophrys (Sandia) macfarlandi 

Due to the considerable interest in this recently discovered “critter,” the following 
previously unpublished records seem worthy of immediate dissemination. Sandia 
macfarlandi has now been recorded in New Mexico from: (a) virtually all points on 
the north, west, and south sides of the Sandia and Manzano Mts. between 5800 and 
6400 ft. in Sandoval, Bernalillo, Valencia, Torrance, and Socorro counties; (b) White 
Oaks and an adjacent colony about 10 miles to the southwest both in Lincoln 
County, and both around 5500 ft. (RH & MT); (c) Alamogordo Lake in De Baca 


County, 4200 ft. (MT); (d) Conchas Lake State Park in San Miguel County, 
4200 ft., and nearby in Guadalupe County on N. M. Route 129 (MT); (e) on the 
eastern side of the Sacramento Mts., near Hondo, Lincoln County (Bruce Harris); 
(£) on the western side of the Sacramento Mts., near High Rolls, 6000 ft., Otero 
County (RH & MT); (g) 3 miles west of Cimarron, 6500 ft., Colfax County (MT); 
(h) on all sides of Ladron Peak around 5800 ft., in Socorro County (RH); (i) on the 
northeast face of the Magdalena Mts., around 6000 ft., in Socorro County (RH); and 
(j) 1 mile NW of Acoma Pueblo, 5500 ft., Valencia County (RH). The last three 
of these records are the only known U. S. occurrences west of the Rio Grande. The 
foodplant of macfarlandi, Nolina texana Wats., does not seem to occur on Mt. 
Taylor or in the San Mateo Mts. Flight season in the Sandias has been found to 
extend from 15 February to 2 July. This makes macfarlandi the first non-hibernator 
to fly in the spring. Average annual temperature low in Albuquerque after 15 
February is 10°F. Under proper conditions, macfarlandi is the most abundant 
butterfly in New Mexico. I have taken 42 with a single swing of a net at the 
composite Senecio longilobus Benth. 


Determinations of Erynnis, Hesperia and Amblyscirtes were made by 
Kilian Roever. John Lane first pointed out that the Sandia and Manzano 
Pholisora populations were not catullus. Chlosyne identifications were 
confirmed by Clifford D. Ferris. Countless hours of discussion about this 
article were spent with Mike Toliver, the only other long-term resident 
Albuquerque collector. 


Beuret, H. 1958. Zur systematischen Stellung einiger wenig bekannter Glauco- 
psychidi. Mitt. entomol. Ges. Basel (N.F.) 8: 61-100. 

Burns, J. M. 1964. Evolution in skipper butterflies of the genus Erynnis. U. 
Calif. Publ. Entomol. 37. iv + 214 p. 

pos Passos, C. F. 1964. A synonymic list of the Nearctic Rhopalocera. Lepid. 
Soc. Mem. 1. vi + 145 p. 

. 1969. A revised synonymic list of the Nearctic Melitaeinae with taxonomic 

notes (Nymphalidae). J. Lepid. Soc. 23: 115-125. 

1970. A revised synonymic catalogue with taxonomic notes on some 
Nearctic Lycaenidae. J. Lepid. Soc. 24: 26-38. 
EMMEL, T. C. 1969. Taxonomy, distribution and biology of the genus Cercyonis 
(Satyridae). I. Characteristics of the genus. J. Lepid. Soc. 23: 165-175. 
Fietp, W. D. 1971. Butterflies of the genus Vanessa and of the resurrected genera 
Bassaris and Cynthia (Lepidoptera: Nymphalidae). Smithsonian Contr. Zool. 
84. 105 p. 

FREEMAN, H. A. 1969. Systematic review of the Megathymidae. J. Lepid. Soc. 
23: Suppl. 1. 59 p. 

Lancston, R. L. 1969. Philotes of North America: synonymic list and distribution 
(Lycaenidae). J. Lepid. Soc. 23: 49-62. 



Department of Entomology, University of California, Davis, California 95616 

The Plebejus acmon group is composed of three closely related butter- 
fly species: P. acmon (Westwood & Hewitson), P. lupini (Boisduval), 
and P. neurona (Skinner) (Goodpasture, 1973). These species com- 
prise the Eriogonum (Polygonaceae )—feeding members of the subgenus 
Icaricia. Plebejus acmon ranges widely in western North America from 
the Pacific Coast west to the Great Plains, P. lupini occurs primarily in 
mountains of the Pacific Coast states, and P. neurona is restricted to ele- 
vations above about 5,000 ft. in central and southern California. Two sub- 
species of P. lupini are recognized; |. lupini occurring north of approxi- 
mately 37° latitude, and l. monticola (Clemence) in southern and Baja 
California. Three subspecies of P. acmon are recognized; a. acmon 
in the southwestern region of the distribution of the species, a. lutzi dos 
Passos in the north, and a. texanus Goodpasture in the south. In Cali- 
fornia, where the distributions of the three species overlap, P. acmon 
is extensively sympatric with P. lupini, but P. neurona, tending to occur 
at higher elevations is only marginally sympatric with P. lupini. 

The purpose of this investigation was to determine the relative degree 
of foodplant specificity among members of the P. acmon group. Data 
suggesting possible biological interactions between foodplant and both 
larval and adult butterflies are discussed. 

Subspecific differences and interspecific variation in foodplant use 
were determined by direct observation in the field. The procedure 
established by Shields et al. (1970) for collecting and accurately report- 
ing foodplant records was followed closely. Observations of larval feeding 
and adult oviposition are recorded in detail in an unpublished M.S. 
thesis (Goodpasture, 1971). Only a brief summary of foodplant records 
is given. 


Larval foodplants of P. neurona. Comstock & Dammers (1933) briefly 
described the early stages of P. neurona and stated that this insect is 
found in association with Eriogonum wrightii Torr. Immature stages they 
described were reared on E. fasciculatum Benth. from, “Eggs secured 
from captive females taken at Blue Ridge, above Wrightwood, San 
Bernardino County on June 8, 1932” (Comstock & Dammers, 1933). 


Subsequent oviposition records and field observations have confirmed 
this association and indicate a definite preference for leaves as oviposition 
sites. Laboratory rearing of adults has been carried out on E. w. var. 
subscaposum ( Wats.) from eggs and first instar larvae found on leaves 
of this plant. 

Foodplant acceptance tests have shown that P. neurona cannot complete 
development on legume plant species that are acceptable to larvae of 
P. acmon acmon (Goodpasture, 1971). 

Evidence of foodplant specificity was obtained from field observations 
made near Mt. Hillyer, Angeles National Forest, Los Angeles Co., 
California, elev. 6,000 ft. At this locality, a number of plants of several 
abundant Eriogonum species were found within a radius of 100 feet. 
Plebejus neurona was the only abundant! Icaricia species encountered 
here during June and July 1969 and 1970 when a systematic search was 
made of the following Eriogonum species at this locality: wrightii var. 
subscaposum, nudum var. publiflorum Benth., davidsonii Greene, and 
umbellatum Torr. ssp. A total of 17 eggs, microscopically identical to 
eggs obtained from captive female P. neurona, were found on E. wrightii 
var. subscaposum, while none were located on the other species of 

Two often sympatric varieties of E. wrightii occur throughout the range 
of P. neurona: E. w. subscaposum in the San Bernardino Mountains north 
into the Sierra Nevada, and E. w. trachygonum (Torr. ex Benth.) from 
northern Los Angeles County to the base of the Sierra Nevada (Munz, 
1968). Foodplant records and field observations indicate that P. newrona 
is restricted to these low, matted shrubs in montane coniferous forest 

Larval foodplants of P. acmon acmon._ Documented records indicate 
that P. acmon acmon feeds on more species of plants than any other 
group member and that it is the only subspecies to utilize legumes as well 
as plants of the family Polygonaceae. 

Localities where observation of utilization of more than one plant 
family as foodplant and or oviposition site have been made are: Monti- 
cello Dam, Napa Co. [Lotus scoparius (Nutt.) Ottley, L. purshianus 
(Benth.), and Erigonum nudum (Dougl. ex Benth.) ], Frazier Park, Kern 
Co. [L. procumbens (Greene) Greene, and Eriogonum sp. (probably 
nudum or elongatum Benth.) ], and Laguna Grade, San Diego Co. [Lotus 
sp. and E. wrightii var. membranaceum Stokes ex Jeps]. At localities 
near Davis, Yolo Co., several foodplants such as L. purshianus and 

1 Plebejus (Icaricia) icarioides (Bdv.) and P. acmon acmon were common at localities within 

at least 1 mi. of the Mt. Hillyer locality, yet none were taken here during the course of this study 
probably due to the absence of appropriate legume foodplants in the immediate Mt. Hillyer area. 

VoLUME 28, NUMBER 1 55 

Polygonum aviculae L., an introduced weed, co-occur with P. acmon 
acmon and are probably utilized simultaneously. At all other localities 
records are available for only one plant species and it is not known if these 
populations are actually or potentially polyphagous. 

At localities such as Monticello Dam where adult flight period extends 
from spring to late summer, suitable foodplants appear to be used 
sequentially according to their seasonal availability. Records from the 
central Coast Ranges in California for Lotus scoparius: 7 March-9 May; 
E. latifolium-nudum: 19 February-15 May (leaves), 3-15 August 
(flowers); and L. purshianus: 21 May-6 September, coincide with 
seasonal availability of these plants. Gorelick (1969) has suggested 
sequential utilization of E. latifolium Sm. in Rees. in the San Bruno 
Mountains, San Mateo Co., with the larvae feeding on leaves in spring 
prior to early summer die-back, and on flowers as they become available 
to successive generations during later months. At Monticello Dam, 
larvae were found to move from drying flower heads of E. nudum 
during late summer to overwintering sites in leaf litter at the base of 
these plants. At other localities, only a single suitable plant species 
may occur as at Putah Creek, U.C.D. campus, Yolo Co., where the only 
foodplant is the annual L. purshianus. Plebejus acmon acmon adults are 
not found at this locality until June (based on two years observation and 
museum specimens), when plants are well established. Seed germination 
of L. purshianus begins very early in winter, with the result that by 
March there are numerous new shoots about four inches long growing 
under dense winter annual vegetation. Progeny of females collected 
at Monticello Dam from March through August and at Putah Creek 
from June through mid-September do not enter diapause when reared 
under laboratory conditions of constant temperature and _ naturally 
occurring daylength. It is assumed that both populations are multivoltine. 
Individuals from these two populations respond similarly to decreasing 
photoperiod and enter diapause in late summer as early instar larvae 
(Goodpasture, 1973). Seasonal flight data indicate that these popu- 
lations differ markedly in response to conditions initiating breaking of 
diapause. Termination of diapause in winter at Putah Creek would seem 
disadvantageous if foodplant is available only in summer months. Ter- 
mination of diapause in winter (December?) at Monticello Dam would 
allow earlier activity of adults and might be advantageous where a po- 
lyphagous population feeds on plants available at different times of the 

A number of plants not known to be fed upon by wild P. acmon larvae 
have been found to be acceptable as laboratory foodplants. These plants 
are Lotus corniculatus L., Lupinis albifrons Benth., Eriogonum fascicu- 


latum, and E. umbellatum (Goodpasture, 1971). In addition, flowers of 
Trifolium obtusiflorum Hook. were reported by Gorelick (1969) to be 
acceptable as larval foodplant. These plants are available to females of 
P. acmon acmon at various localities but are apparently not selected 
for oviposition and are probably not utilized as food. Eriogonum 
fasciculatum, for example, is coextensive with P. aemon acmon in southern 
California, but appears not to be used as food. 

At many localities the shrub-like E. fasciculatum occurs and other, 
usually herbaceous species of Eriogonum and/or legumes are used as 
foodplants. For example, at Del Puerto Canyon, Stanislaus Co., and 
Frazier Park, E. nudum is the only known foodplant. Field observations 
of foodplant utilization, and adult flight period, and Eriogonum ecology, 
show that at these localities E. fasciculatum is potentially available to 
larvae and ovipositing females from March or April through June. At 
other localities such as Switzer’s Camp, Los Angeles Co., and Laguna 
Grade, adult flight season extends from June through September or 
October when E. fasciculatum is probably not suitable as larval food- 
plant due to cessation of vegetative growth of this plant during summer 

Preference for oviposition on certain plant parts may exist in some 
populations. At some localities, oviposition has been observed on all 
plant parts (Lotus, Switzer’s Camp and Putah Creek), exclusively on 
leaves (Eriogonum wrightii, Laguna Grade), or exclusively on flowers 
(E. elongatum, Hidden Valley and Lake Sherwood, Ventura Co.). Where 
Eriogonum species are utilized as foodplant, eggs might be placed on 
floral or leaf structures depending on seasonal availability or suitability 
of plant material. 

In conclusion, the foodplants of P. acmon acmon in California are 
plants of the families Leguminosae and Polygonaceae. As can be seen 
in Fig. 1, the perennial herbaceous Eriogonum species latifolium, nudum, 
and elongatum, as well as certain legumes (Lotus scoparius and L. 
purshianus ), are the most frequently encountered foodplants. In south- 
ern California, Lotus species may serve as the primary food source, 
with several additional Eriogonum species of rather limited distribution 
(e.g. E. parvifolium Sm. in Rees. and E. plumatella Dur. & Hilg.) as 
occasional foodplants. 

Larval foodplants of P. acmon lutzi and P. acmon texanus. Foodplants 
of P. acmon occurring outside of California are poorly known. Available 
records are primarily associational and indicate that various Eriogonum 
species are utilized as foodplants. 

Association of adults of P. acmon lutzi with E. marifolium T. & G. and 

VoLUME 28, NUMBER 1 57 

E. pyroliifolium Hook at Mt. Bachelor, Oregon suggests use of these 
plants. Adults of P. a. lutzi form spangelatus have been reared from 
larvae collected on an unidentified Eriogonum species, Olympic Moun- 
tains National Park, Washington (J. Pelham, pers. comm.). Foodplant 
records for Wyoming include E. flavum Nutt. and E. umbellatum (C. D. 
Ferris, pers. comm.). 

Observations of oviposition and co-occurrence of adults and plants 
suggest that P. acmon texanus feeds on E. wrightii var. wrightii at lo- 
calities in Arizona, New Mexico, and Texas, and that E. corymbosum var. 
velututinum Reveal & Brotherson is a foodplant near Cerrillos, New 
Mexico. Oviposition records, as well as associational data, indicate that 
E. effusum Nutt. and E. racemosum Nutt. are utilized at several Colorado 

Larval foodplants of P. lupini lupini. Several shrub-like Eriogonum 
species are documented as larval foodplants of P. lupini lupini. In Cali- 
fornia, E. umbellatum and E. ovalifolium Nutt. are the only known food- 
plants. Emmel & Emmel (1962); and Garth & Tilden (1963) have noted 
adult association with Eriogonum at Donner Pass and in Yosemite 
National Park. It appears from available records, that E. umbellatum 
is the primary food source in the Sierra Nevada and the north Coast 
Ranges at least in California. In Nevada, a wider variety of Eriogonum 
species may be utilized. Records from mountainous areas in central 
Nevada indicate use of E. kearneyi Tidestr., E. ovalifolium, E. pal- 
merianum Reveal, and E. umbellatum. Statements that Lupinus spp. 
are utilized ( Boisduval, 1869, “. . . dans le sud de la Californie.”; Jones, 
1951, Washington) are without supporting data. 

Field observations made at Echo Lake, E] Dorado Co., suggest that 
flowers of E. umbellatum var. umbellatum are preferred as an ovi- 
positional site. Seven females seen ovipositing at this locality (13-19 
July 1970) laid a total of 17 eggs, 15 on flowers and two on leaves. 
Females were not seen to oviposit on other Eriogonum species (nudum, 
incanum Torr. & Gray, lobii Torr. & Gray, and wrightii) also present 
at this locality. 

Plebejus lupini appears most similar to P. neurona in terms of larval 
acceptance and the nutritional adequacy of several plants, as well as in 
preferences inferred from field data. Larvae of these two species accept 
flowers of Lotus corniculatus but develop poorly and suffer high mor- 
tality, whereas P. acmon acmon larvae show no mortality on this plant 
(Goodpasture, 1971). 

Larval foodplants of P. lupini monticola. Foodplant records from eight 
localities suggest that P. lupini monticola is restricted to E. fasciculatum 


” cs 
— B. = 
= =. = = = 
Ee aa =< ves = 
w cs _ ae Ee 
ms = = i— +. 
= Ee = Ee = = 
co oO o o s = 
i co os os — — 
Foodplant Pee Cee 

Polygonum aviculae ---- - ---- 

Eriogonum corymbosum- --- -- 
Bit (UiSsLT eee ae are 
elloneattiimes == ee 
fascrculatiim= 22222552 S2 
kearneyi----- 2 
latifolium & nudum - ---- -- 
marifolium & pyroliifolium - - 
OVA ONIN = 25252 = 2 4: 
NAME WNW sae a 
paVitOtiMess === 22 oes 
Dlumatelliae= === = = 2 
PacCeMaSumes =] oa = 
Umibye Hatem 
QWWCMEIMM SSe=2===2= == 
WiTiCiitelem Wit SONIC i 
w. SubScaposum & 2-22 — 

w. trachygonum 

Lotus & Astragalus --- ---- 

Fig. 1. Summary of foodplant records for the Plebejus acmon group. Docu- 
mented records (D) = larval rearing to adult, oviposition observed. Suspected 
foodplants (S) = adults collected in association with plant, pre-oviposition behavior 
observed. Numerals refer to number of localities (more than 10 miles apart) 
where observations of foodplant use have been made. 

in most of its range throughout chapparral communities in the southern 
half of California. Utilization of more than one species of Eriogonum 
may occur at Laguna Grade, where females have been observed to 
oviposit on E. wrightii var. membranaceum as well as on E. f. var. 
polifolium (Benth.). At this locality, near the upper altitudinal limit 
of E. fasciculatum, E. w. var. membranaceum is the more common plant, 




cteioontene Okie 

Fig. 2. Geographical distribution of foodplant specificity in Plebejus acmon. 
Dashed line represents approximate known distribution of P. acmon. Dotted line 
represents distribution of P. lupini and area of P. acmon—P. lupini sympatry. P. lupini 
foodplants are shrub-like Eriogonum species throughout its range. A = Astragalus, 
L = Lupinus, P = Polygonum, HE = herbaceous Eriogonum, SE = shrub-like 
Eriogonum species. Herbaceous Eriogonum species include elongatum, latifolium, 

and nudum. Other species of Eriogonum mentioned in the text are considered shrub- 
like (Munz, 1968). 


and appears to be the principal foodplant. Oviposition by a single female 
has been observed on E. umbellatum subsp. at Horse Thief Springs, 
San Bernardino Co. Other Eriogonum species present at this locality 
include fasciculatum, wrightii, and heermannii Dur. & Hilg. (A. O. 
Shields, pers. comm.) and may also be utilized by P. lupini monticola. 

Observations of oviposition behavior and placement of field collected 
eggs indicate that floral structures are preferred over leaves as oviposition 
sites when both are available (Goodpasture, 1971). 

Geographical distribution of foodplant specificity. Fig. 1 illustrates 
that members of the Plebejus acmon group tend to have mutually ex- 
clusive diets. This is especially evident in California where distributions 
of four of these entities overlap. Plebejus neurona, P. lupini monticola, 
and P. lupini lupini have no foodplants in common at the varietal level 
and are narrowly oligophagous, feeding predominantly on a single species 
or subspecies of Eriogonum. Outside of California, at least in central 
Nevada, P. lupini apparently utilizes a much wider variety of Eriogonum 
foodplants. As can be seen in Fig. 2, polyphagy in P. acmon shows some 
correlation with geography and distribution of P. lupini. Thus, P. acmon 
feeds on Astragalus (A), Lotus (L), Polygonum (P), herbaceous (HE), 
and shrub-like Eriogonum (SE) species only in California where it is 
broadly sympatric with P. lupini. Outside of California, P. acmon is 
known to utilize only shrub-like Eriogonum species. 


Larval choice of foodplant in Lepidoptera may be requisite for survival 
in species with a larval dispersal stage or with larvae defoliating part of 
their available food supply (Dethier, 1959; Cook, 1961). Larval ability 
to select proper plant species may also play a role in foodplant relation- 
ships in this Plebejus group where larvae overwinter as early instars. 
For example, P. acmon acmon terminating diapause in May at Putah 
Creek must locate early season growth of the annual Lotus purshianus. 
Larvae terminating diapause at localities where annual foodplants are 
utilized may encounter and accept plant species other than those upon 
which eggs were laid during the previous growing season. The prob- 
ability of larvae encountering foodplants of the same species as chosen 
by females of the final summer generation may depend on such factors 
as success of seedling establishment in annual plants or the number of 
acceptable alternative plants growing in close proximity to overwintering 

Larval acceptance tests have demonstrated that larvae from several 
populations of several subspecies of the P. acmon group accept many 
plant species not known to be utilized in nature. While these potential 


foodplants elicit and sustain, or at least do not deter, larval feeding, full 
suitability? remains uncertain because fertility of reared adults has not 
been tested. It is assumed from larval acceptance of various plants, as 
well as from knowledge of plant distributions, that a large number of 
Eriogonum species are available and suitable to many populations of all 
group entities. At least some populations of P. lupini and P. neurona feed 
on only one of several species of Eriogonum growing at one locality. A 
similar pattern of food resource utilization may exist throughout the 
Icaricia. It has been shown that larvae of P. icarioides will feed on any 
species of Lupinus in captivity, but wild populations normally utilize only 
a few of the possible range of Lupinus species growing locally (Downey 
& Fuller, 1961; Downey, 1962). 

Rigid specificity encountered in nature in most members of this 
group seems to be due primarily to the precision with which females lay 
their eggs on certain plants, as has been stressed by Merz (1959) for 
several other Lepidoptera. In field studies, both P. neurona and P. 
lupini lupini were found to display a high degree of foodplant specificity. 
Presumably, females respond to specific plant stimuli in selecting a 
single Eriogonum species for oviposition from among several available 
at a single locality. 

Many populations of the P. acmon group differ in both plant species 
and complement of species utilized for food. Although this may be the 
result of coincidence of plant distribution, larvae and ovipositing females 
may have different preferences at different localities. It should be 
noted that the extent to which differences in foodplant use reflect 
differences in foodplant preference is unknown in this group. 

Failure of P. acmon members to utilize certain potential foodplant 
species during certain times of the year, e.g. E. fasciculatum by P. 
acmon acmon during summer months, may be due to unsuitability caused 
by drying and hardening of leaves and flowers. Although females of 
at least some P. acmon acmon populations will lay eggs on E. fasciculatum 
var. foliolosum in captivity, larvae are not able to survive on this plant 
when hatching from eggs laid during and after June (Goodpasture, 
1971). Cole (1967) has shown that shoot growth of several Eriogonum 
species, including E. fasciculatum, ceases in May in the Santa Monica 
Mountains and that dramatic changes in leaf physiology accompany soil 
drought in summer months. 

Differences in feeding preferences within and between closely related 
species may provide information on evolutionary mechanisms that can 

2 Remington & Pease (1955) define the test of full suitability of a plant in terms of larval 

rearing to adult solely on that plant with the production of adults which, when induced to 
mate, lay eggs which then hatch. 


account for changes between polyphagy and monophagy ( Dethier, 1954). 
For example, within this group spatial and temporal differences in 
foodplant utilization may have evolved independently in isolation or as a 
result of competition resulting in ecological character displacement. The 
data available do not allow distinction between these alternatives. How- 
ever, evidence seemingly in support of competitive displacement is: 
(1) different sympatric subspecies of this group tend to have mutually 
exclusive foodplants; (2) P. acmon does not utilize shrub-like Eriogonum 
species where it is sympatric with P. lupini, and feeds on a wide variety 
of shrub-like Eriogonum species where it is widely allopatric to P. 
lupini; and (3) P. acmon is morphologically distinct from P. lupini 
where these two species are sympatric, and convergent to P. lupini 
where it occurs in states widely allopatric to the distribution of P. lupini 
(Goodpasture, 1973). 

Dethier (1954) has also suggested that polyphagy is the more primitive 
condition in phytophagous insects. This may also apply to the P. acmon 
group. Oligophagy and Eriogonum feeding as exemplified by the narrow 
feeding habits of P. neurona may have been derived from a polyphagous 
ancestor with food habits similar to those of P. acmon acmon. 


Data on foodplants use by members of the Plebejus acmon group are 
summarized in Fig. 1. Plebejus acmon acmon is the only polyphagous 
group member, feeding on the legumes Lotus and Astragalus as well as 
the Polygonaceae Eriogonum and Polygonum. All other group members 
are oligophagous and restricted to feeding on one or a few species of 
Eriogonum. Members of this group tend to have mutually exclusive diets. 

Differences in foodplant use between species and subspecies of this 
group are discussed in terms of foodplant ecology, geographical distribu- 
tion of foodplant specificity, and possible larval feeding and adult 
oviposition preferences. 


Many of the foodplant records used in this study were collected by 
Oakley Shields (University of California at Davis) and John Emmel 
(Santa Monica, California), and their assistance is gratefully acknowl- 
edged. Acknowledgment is also due Chris Henne (Pearblossom, Cali- 
fornia), Paul Opler, James Scott, John Shepard (formerly University of 
California at Berkeley), R. F. Denno, A. M. Shapiro ( University of Cali- 
fornia, Davis), C. D. Ferris (University of Wyoming), and John Lane 
(California State University at Northridge) for providing valuable field 
data and making collections available for study. 

VOLUME 28, NuMBER 1 63 

Numerous plant determinations were kindly made by Dr. J. L. Reveal 
(University of Maryland) for Eriogonum, B. J. McCaskill ( University 
of California, Davis), and R. Gustafson (Los Angeles County Museum 
of Natural History). 

For their aid in discussion and constructive criticism throughout this 
study I am indebted to Drs. R. W. Thorp, R. M. Bohart, L. D. Gottleib, 
and Mr. Oakley Shields. 


BotspuvaL, J. A. 1869. Lepidopteres de la Californie. Ann. Soc. Entomol. Belg. 
12: 8-94. 

CoE, N. H. A. 1967. Comparative physiological ecology of the genus Eriogonum 
in the Santa Monica Mountains, Southern California. Ecol. Monogr. 37: 1-24. 

Comstock, J. A. & C. M. Dammers. 1933. Notes on the life histories of four 
Californian Lepidopterous insects. Bull. So. Calif. Acad. Sci. 32: 77-83. 

Coox, L. M. 1961. Foodplant specialization in the moth Panaxia dominula L. 
Evolution 15: 478—485. 

Deruier, V. G. 1954. Evolution of feeding preferences in phytophagous insects. 
Evolution 8: 33-54. 

1959. Foodplant distribution and density and larval dispersal as factors 
affecting insect populations. Can. Entomol. 91: 581-596. 

Downey, J. C. 1962. MHostplant relations as data for butterfly classification. Syst. 
Zool. 11: 150-159. 

. & W. C. Futter. 1961. Variation in Plebejus icarioides (Lycaenidae). I. 
Foodplant specificity. J. Lepid. Soc. 15: 34-42. 

EMMEL, T. C. & J. F. EMMEL. 1962. Ecological studies of Rhopalocera in a high 
Sierran community—Donner Pass, California. I. Butterfly associations and dis- 
tributional factors. J. Lepid. Soc. 16: 23-44. 

GartH, J. S. & J. W. TitpEN. 1963. Yosemite butterflies. J. Res. Lepid. 2: 1-96. 

GooppasturE, C. E. 1971. Biology and systematics of the Plebejus (Icaricia) 
acmon group. Unpub. M.S. thesis, Univ. Calif., Davis. 

1973. Biology and systematics of the Plebejus (Icaricia) acmon group. I. 
Review of the group. J. Kansas Entomol. Soc. 46: 468-485. 

Goretick, G. A. 1969. Notes on larval host acceptance in a California population 
of Plebejus acmon. J. Lepid. Soc. 23: 31-32. 

Jones, J. R. J. L. 1951. An amnotated check list of the Macrolepidoptera of 
British Columbia. Entomol. Soc. B.C., Occ. Paper No. 1: 1-148. 

Merz, E. 1959. Pflanzen und Raupen. Uber einige Prinzipien der Futterwahl bei 
Goss-schmetterlingsraupen. Biol. Zentr. 78: 152-188. 

Munz, P. A. 1968. Supplement to a California flora. Univ. Calif. Press, Berkeley 
and Los Angeles, 224 p. 

ReMinctTon, C. L. & R. W. PEAsE, JR. 1955. Studies in foodplant specificity. I. 
The suitability of Swamp White Cedar for Mitoura gryneus (Lycaenidae). 
Lepid. News 9: 4-6. 

SHIELDs, O., J. F. Emmet & D. E. Breeptove. 1970. Butterfly larval foodplant 
records and’ a procedure for reporting foodplants. J. Res. Lepid. 8: 21-36. 



60 Estes Street, Lakewood, Colorado 80226 

The purpose of this paper is to describe adult behavior (mate- 
locating, mating, feeding, oviposition, and basking), dispersal, and popu- 
lation parameters (especially lifespan) of L. arota Boisduval. 

A mark-recapture study was conducted in 1969 at the mouth of Rouch 
Gulch (Spring Creek), Fremont County, Colorado. L. arota was also 
studied in 1971 one km east of Smith Creek Campground, Custer County, 
Colorado, and at Little Fountain Creek, E] Paso County, Colorado. These 
sites were mountain gulches or streams with abundant larval host, 
Ribes spp. 


Number of matings per female was determined by counting spermato- 
phores (Burns, 1968). For study of movements and estimation of popu- 
lation parameters, butterflies were marked individually using the method 
of Ehrlich & Davidson (1960). They were captured with a net, marked, 
and immediately released at the site of capture. 

Analysis of population movements. The following method allows direct 
comparison between the sexes and between species, determination of 
change of movements with age, and separation of the velocity and 
distance aspects of movements. On a map of the movements of each 
recaptured individual, distances in mm between each two captures are 
measured, and called d, between first and second captures, d. between 
second and third, etc. Total distance moved by the individual, D, is the 
sum of the d’s. Range, R, an estimate of dispersal, is distance between 
the two most distant capture points. Time in days between first and 
second capture is called t,, etc. T is total time between first and last 
capture. Velocities are defined v; = d;/t; and V = D/T. Midpoint age 
between captures is determined by finding the age midway between two 
captures after calling the first capture day 0. Correlations between 
distance or velocity and midpoint age determine whether movements 
change with age. Midpoint age is used rather than age at start or end 
of a period between captures because the time between captures differs. 

Jolly’s stochastic method was used to estimate population size, survival 
rates, and number of new individuals joining the population (Jolly, 
1966). The method of Cook et al. (1967) was used to obtain expected 
lifespan from average survival rate. 

VoLUME 28, NUMBER 1 65 


Mate-locating behavior. To locate females, males perch on branches 
of shrubs and trees 1 to 2 m above the ground. Males start perching at 
about 0715 and actively perch until 1100. Males gradually stop perching 
between 1100 and 1200, and rarely perch after 1200. In the afternoon, 
males mainly visit flowers, or are quiescent on shrubs. Perching males 
sit on a branch or leaf and dart out at passing objects. Most such objects 
are other males, whereupon the two males fly about each other for a 
few seconds, then each returns to or near the same perch as before. 
One male was observed to perch on a one-meter length of oak branch 
for two hours during which time he made almost 100 short flights at 
passing objects. Other males sometimes flew short distances between 
investigative flights. Males chase a narrow size range of passing objects, 
so that almost all of the chases are toward other L. arota. 

Males perch in small clearings in many different topographic situations. 
A clearing likely to have perching males is about 3-5 m. in diameter, rea- 
sonably level, and surrounded by tall trees or steep hills or both. Males 
were observed to perch in clearings that were in a small valley bottom, 
on a hillside along an irrigation ditch, and on a ridgetop. More males 
perch in valleys than hillsides or hilltops because more suitable clear- 
ings are found there and the larval hostplant is usually more abundant 

Mating. If a female flies past a perched male, the male darts after 
her and flies about 14 cm below her for several meters, then the female 
lands on a branch and the male lands behind. The male then usually 
flicks his wings by holding them about 60° from the vertical (about 120° 
from each other) and vibrating them at small amplitude (only 1-2 
mm amplitude for each wing). Sometimes the male vibrates the wings 
only once or twice per second, and other times he vibrates his wings 
rapidly (about 10 times per second), occasionally with the vibrations 
clustered into groups. These two types of wing flicking occurred about 
equally often, but only one type was seen in any one courtship, except 
for one courtship in which the male vibrated his wings once per second, 
then drew closer to the female and vibrated his wings about 10 times per 
second. Other males did not flick their wings at all, but merely walked 
to the female and attempted copulation. Whether or not the male 
flicks his wings, if the female remains quiescent the male crawls along- 
side and bends his abdomen either right or left to attempt copulation. 
In the two completed copulations observed, the female was quiescent 
during courtship and mating. Unreceptive females flap the wings almost 
full stroke about 10 or more times per second for about 2-5 seconds, while 


TaBLE 1, Dispersal data for Lycaena arota. N = sample size. Dispersal param- 
eters are defined in methods section. 

Dispersal Parameter Males N Females N 
Number Marked 107 94 

Number Recaptured 53 ot 

Average T (days) 2.56 5S 1.91 37 
Average ti (days) yl ala! 84 85 
Average R (meters) 15 53 29 Sih 
Average D (meters) 16 53 29 Oi 
Average di (meters) 6 144 13 85 
Average V (meters per day) 1a 53 18 oy 
Average vi (meters per day) 5 144 9 85 

sitting. In most courtships the female was previously mated, and she 
performed this “rejection dance” when the male crawled up to her; 
the male flew away then or after subsequent rejection dances. Male wing 
flicking may cause the female to become quiescent and receptive, because 
if the female is already quiescent, the male usually does not flick his wings. 

Courtship and mating occur at the same time as male perching. 
Copulating pairs were found at 0805, 0920, 1121 and 1200, all in valley 
bottoms by Scott, and at 0930 and 1326 (Oakley Shields, pers. comm.). 
1326 is after the normal perching period, and that observation may 
represent the mating of a late perching male, or perhaps the mating was 
initiated during the normal perching period and the pair remained joined 
until observed (copulation of butterflies lasts rarely up to 30 hours). 
Nineteen courtships were observed from 0815 to 1036, and one was 
observed at 1121. 

Females rarely mate more than once. Of 60 females dissected (caught 
several weeks after the species had first appeared), 15 were virgin, 44 
had mated once, and only 1 had mated twice. Many virgin females were 
found in the afternoon, indicating that many females wait until the 
day after emergence to mate. 

Movements. The mark-recapture study was carried out from 30 July 
to § August 1969. Rouch Gulch is a small, dry (except after rains) gulch 
opening into the Arkansas River, within the pinyon-juniper belt. The 
larval host, Ribes leptanthum, and adult nectar sources were scattered 
along the bottom, where the recapture study was carried out from the 
Arkansas River to 300 m. up the gulch. Nearby gulches were sampled 
to detect dispersal. The proportion of recapture (Table 1) was higher 
for males, probably because males disperse less than females. Although 



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V9r + 189 6ST’ + POT’ 9'F9IT + 6602 0¢S'08 968c° S 
c06 +916 Sc)’ + 166’ Voy + L Sct Rey, gece’ [ Jsusny 
v69 + T&T Ca COOI GiGi lene Ol Sl SP 18SG" Ig 
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‘1019 plepue}s—yS ‘uorjepndod oy} suturol 
sjewiue Mou JO oquInu—g ‘[eATAIns Jo AyTIqeqo1d—iyq ‘uorjeindod [e}0}—N ‘uonejndod poxreu [e}0}—] ‘sjeulue pexieul fo uor}10d 
-o1d—vydiy “(9961 ) AT[O[ JO Jopourt o1jseyooys oy} SuIsn veep ainjdeooer o[dy[nu woIF pozeutsjse stojourvied uonjeindog °% ATaV I, 


TABLE 3. Flowers and other fluid sources visited by L. arota at the three study 

Little Gulch Smith 
Fountain ————_—__————— Creek 
Species Color Creek Male Female Cmpgd. 

Solidago occidentalis yellow few 236 158 
Pericome caudata yellow 52 18 
Eriogonum jamesi whitish-yellow few 30 64 
Heterotheca villosa yellow 5) 9 
Chrysothamnus nauseosus yellow 6 5 
Helianthus pumilus yellow 2 2 
Allium sp. white 1 
Rubus parviflorus* — 5 3 
mud — | 
Rudbeckia laciniata yellow many 
Melilotus alba white many 
Aster novae-angliae bluish white many 
Apocynum sp. white many 
Clematis sp. cream white few common 
Achillea sp. white few 

1 Juices of the blue-black berries. 

the times between captures for females were less than those for males, 
the distances moved were much greater for females, resulting in female 
velocities almost twice those of males. Most individuals of both sexes 
did not move at all between recaptures, so that the averages of movement 
statistics in Table 1 are low. Some individuals move considerable dis- 
tances, however. Maximum distances moved (ranges) for males were 
83, 84, 92, 93, 168, and 214 meters, and for females 82, 83, 85, 92, 94, 
169, and 186 meters. 

Population parameters. The population size in the study area at Rouch 
Gulch, which is about 300 m. by 100 m., was about 400, declining to 200 
at the study’s end (Table 2). Females were slightly more common than 
males since males emerge several days before females and the mark- 
recapture study was conducted after the peak emergence of males. The 
number of new animals joining the population was fairly high, about 
60 per day for males and 80 for females. The number of animals joining 
the population and the survival rates are due mostly to emergence 
and deaths respectively, because few individuals dispersed out or into 
the area. The 0.164 survival rate for males is low because it represents 
the survival over a three day period. The average survival rate for 
males and corresponding expected lifespan was 0.763 (3.7 days) using 

VoLUME 28, NUMBER 1 69 



OF 7© 


ie 30 



Fig. 1. Number of visits to flowers at various times of day for both sexes of L. arota. 

method 1, and 0.752 (3.5 days) using method 2 (methods of Scott, 
1973). The rate for females was 0.790 (4.2 days) using method 1 and 
0.725 (3.1 days) using method 2. The potential lifespan is much longer: 
males have lived at least 8, and females at least 6, days. The survival 
rates for both sexes were undoubtedly decreased by extensive predation 
by robberflies and ambush bugs (see below) and by the weather, which 
was very hot especially toward the end of the study when the survival 
rate declined slightly. 

Feeding. Both sexes very often feed at flowers (Fig. 1). Flower visits 
were recorded for 336 males and 296 females. Before 1100, when males 
are perching, males visit flowers less than females, but after 1100 males 
visit flowers more than females: 32 males and 58 females were recorded 
before 1100; 304 males and 238 females after 1100. Both sexes prefer 
yellow and white flowers but feed on different species at different locali- 
ties (Table 3). Males preferred Solidago occidentalis at Rouch Gulch 
slightly more than females, and females preferred Pericome caudata 


and Eriogonum jamesi more than males, but this difference seemed 
mainly because Solidago was concentrated at the mouth of the gulch 
(where males were more abundant) whereas the other plants were more 
widely distributed. The preference for yellow flowers provided ambush 
bugs with many meals (see predation below). 

Oviposition. Females oviposit on twigs, bark, or dead leaves, on or 
under bushes of Ribes species, especially the subgenus Grossularia. 
Many ovipositions on R. leptanthum were observed at Rouch Gulch and 
Little Fountain Creek. Two ovipositions on R. montigenum were ob- 
served at Williams Canyon, E] Paso County, Colorado. John Emmel 
(pers. comm.) found ova on R. leptanthum near Glenwood Springs, 
Garfield Co., Colorado. Emmel et al. (1970) found larvae on R. 
velutinum in Nevada which were raised to adults. Gunder (1930) found 
larvae on Ribes gracillimum in southern California. Females often spend 
an hour or more on the same or a nearby Ribes bush alternating oviposi- 
tion about every five minutes with basking, “hindwing rubbing,” and 
sitting. Females oviposit in the center of a bush just as often as on the 
outer branches. Nine eggs were laid on rough bark of the thicker 
branches, 12 were laid on sides of smooth twigs of the thinner branches, 
and 2 were laid on dead leaves of two different dicotyledons underneath 
Ribes bushes. Females oviposit during the warmer hours of the day 
from 0900 to at least 1430. The eggs undergo diapause and do not hatch 
until the next year. 

Thermoregulation. Both sexes bask by holding the wings about 60° 
from vertical, 120° from each other, the same position as in male courtship 
flicking. Individuals orient their body in any position which brings the 
Wings somewhat perpendicular to the sun, but usually face away from 
the sun. Both sexes bask more often in morning and late afternoon. A 
few individuals basked while feeding, but basking is more frequent 
between oviposition and “hindwing rubbing.” 

Roosting. One male was found on the leaf of an oak tree (Quercus 
gambellii) at 1708, at Smith Creek Campground, Custer County, 

Predation. Considerable predation was observed. Robberflies (Asil- 
idae) captured 4 males and 2 females, and missed others. Ambush bugs 
(Phymatidae) caught one male on Solidago flowers and 1 male and 3 
females on Pericome flowers. A crab spider (Thomisidae) on a composite 
species caught a female at Little Fountain Creek. 

L. arota (and all other Lycaenidae except Riodininae) move the hind- 
wings forward and back, the left and right wing moving in opposite 
directions, a behavior which I call “hindwing rubbing.” Many males 

VoLUME 28, NUMBER 1] iq 

and females did this at all times of day. Basking and hindwing rubbing 
often follow each other, but do not occur often at flowers. L. arota has 
2 mm tails on the hindwing, but does not have a conspicuous eyespot 
next to the tail, so hindwing rubbing may or may not serve to divert bird 
attack away from the body by drawing attention to the antenna-like tail, 
as hypothesized for other Lycaenidae (see Wickler, 1968). One male 
and two females had both hindwings truncated in the manner which may 
indicate bird or lizard attack. 


L. arota differs from L. xanthoides Boisduval and many other Lycaena 
in that mating occurs only in early morning. The preference of L. arota 
for perching in small clearings contrasts with the open perching sites 
chosen by L. xanthoides; these perching sites correspond to the usual 
habitats of the two species, mountain foothills for L. arota and flat 
open areas for L. xanthoides. Population movements of L. arota are 
small, but several factors combine to counteract the weak dispersal: 
(1) females disperse much more than males because their velocities 
are greater and they live longer; (2) the dispersal of some individuals 
is much greater than that of most others; (3) larval hostplants are 
rather scattered, resulting in more continuous populations than if the 
plants were aggregated and these aggregations widely separated. L. 
arota is found almost everywhere in southern Colorado at least between 
1500 and 2200 m. in the Wet Mountains and Arkansas River Canyon; 
and (4) the species is usually very abundant, so that more individuals 
will move over long distances than if the species was less abundant. 
Small peripheral populations could therefore be swamped by a few 
immigrants from a larger population. The species has four named sub- 
species, but the amount of geographic variation is small considering the 
size of the range and the many geographic barriers therein. Courtship 
is very similar among several species of Lycaena. Wing flapping of 
females seems to be a rejection dance in L. xanthoides, L. gorgon 
Boisduval, and L. helloides Boisduval. L. xanthoides males flap the 
wings with wide amplitude and hover behind or over the female, rarely 
flapping while sitting behind her. The courtship of L. gorgon appears 
identical to that of L. xanthoides, and the courtship of L. helloides is 
very similar to that of L. xanthoides except that male helloides vibrate the 
wings with slightly less amplitude while holding them about 30° above 
horizontal. Other aspects of behavior which are similar in L. arota and 
L. xanthoides include basking, feeding behavior (L. arota may be more 
restricted to yellow flowers), and “hindwing rubbing.” 



Males perch from 0715 to 1100 on shrubs or trees in small clearings and 
dart out at passing objects in search of females. Mating usually involves 
male wing flicking. Unreceptive females flutter their wings until the male 
departs. Adults are very sedentary, although some individuals move 
several hundred meters. Both sexes live an average of only 4 days, 
probably because of hot weather and extensive predation. Both sexes 
often feed, usually on yellow or white flowers. Females oviposit mainly 
on Ribes branches. 


I thank Jerry A. Powell for improving the manuscript. 


Burns, J. M. 1968. Mating frequency in natural populations of skippers and 
butterflies as determined by spermatophore counts. Proc. Nat. Acad. Sci., 
Washington, D.C. 61: 852-859. 

Cook, L. M., L. P. BRowEer & H. J. Croze. 1967. The accuracy of a population 
estimation from multiple recapture data. J. Anim. Ecol. 36: 57-60. 

EnruicH, P. R. & S. E. Daviwson. 1960. Techniques for capture-recapture 
studies of Lepidoptera populations. J. Lepid. Soc. 14: 227-230. 

EMMEL, J. F., O. SHiELDs & D. E. BREEDLOVE. 1970. Larval foodplant records for 
North American rhopalocera. Part 2. J. Res. Lepid. 9: 233-242. 

Gunper, J. D. 1930. Butterflies of Los Angeles County, California. Bull. So. 
Calif. Acad. Sci. 29: 39-95. 

Jotty, G. M. 1966. Explicit estimates from capture-recapture data with both 
death and immigration—stochastic model. Biometrika 52: 225-247. 

Scott, J. A. 1973. Convergence of population biology and adult behaviour in two 
sympatric butterflies, Neominois ridingsii (Papilionoidea, Nymphalidae) and 
Amblyscirtes simius (Hesperioidea, Hesperiidae). J. Anim. Ecol. (in press). 

Wicker, W. 1968. Mimicry in Plants and Animals. McGraw-Hill, New York. 
160 p. 


On 15 October 1972, a wild male interspecific hybrid between Limenitis arthemis 
astyanax (Fabricius) and L. archippus (Cramer) [form rubidus Strecker] was 
captured in Durham County nine miles south of Durham, North Carolina. This 
specimen was captured approximately one-half mile from the site at which a 
similar hybrid was found two years previously on 10 October 1970 (Platt & Green- 
field 1971, J. Lepid. Soc. 25: 278-284). The specimen presently is in the collection 
of A. P. Platt at the University of Maryland Baltimore County. 

The recent hybrid (Fig. 1) is in good condition, showing little evidence of being 
flight-worn. It was quite vigorous in flight and eluded capture several times before 
being netted. This specimen closely resembles the lab-reared dark morph depicted 
by Platt & Greenfield (1971); it entirely lacks all remnants of the medial partial 
white banding (an archippus character). On the other hand, the previously caught 
wild hybrid from Durham was battered, and clearly showed traces of the white 

VoLUME 28, NuMBER 1 73 

Fig. 1. Wild-caught male L. arthemis astyanax x L. archippus (hybrid form 

rubidus Stkr.) collected south of Durham, North Carolina on 15 October 1972 by 
J. C. Greenfield, Jr. Above: dorsal surface; below: ventral surface. 


banding on the ventrum of the forewings. Otherwise, the two specimens appear to be 
morphologically identical, both representing the dark morph of rubidus. 

The purpose of this note, in addition to reporting the new hybrid specimen, is to 
consider why two species as morphologically dissimilar in color pattern as the 
red-spotted purple and the viceroy should engage in interspecific hybridization in 
nature, even to the limited extent indicated by the capture of these rare rubidus 
hybrids. The occurrence of two (which are presumably genetically unrelated) in the 
same locality two years apart is unlikely in view of the scarcity of this form in 
the wild. A simple explanation is not readily forthcoming. 

Adults of the two parental species are, however, quite prevalent in this area; both 
astyanax and archippus have been encountered in the same vicinity on multiple 
occasions, especially in the late summer, often congregating in an old orchard where 
the ground is covered with rotting fruit. Thus, the usual ecological separation of 
the two parental species by habitat preference does not seem to be as strongly in 
evidence in this area as in others. 

The fact that both hybrids were collected at almost the identical time of year 
suggests that there exists a greater tendency for natural hybridization in the late 
summer and early fall. This contention may be substantiated by examining the 
collection dates for the known wild Limenitis hybrids listed by Platt & Greenfield 
(1971). Of ten specimens for which dates are reported, nine (including all speci- 
mens of form rubidus) were collected in the months of August and September. 

The single exception to this generality represents the most northern record listed: 
a hybrid specimen presumed to represent a cross between the western subspecies, 
L. arthemis rubrofasciata Barnes & McDunnough, and L. archippus, collected at 
Beulah, Manitoba on 29 June 1904 by A. J. Dennis (Gunder 1934, Canad. Entomol. 
66: 39-48). Although originally described under the hybrid name rubrofascechippus 
(Gunder), it clearly represents a local variant of hybrid form arthechippus (Scudder), 
and has been so listed by Platt & Greenfield (1971). Both the northern locality 
and the early collection date suggest that it also resulted from a fall mating, since 
the larva would undoubtedly have had to overwinter in order to produce an adult by 
late June. Finally, the most recent record of a rather aberrant morph of arthechippus 
(Johnson & Malick 1972, Rep. Flora & Fauna Wisc. 7: 1-6) also has an August 
collection date. 

Both L. a. astyanax and L. archippus are at least partially triple brooded through- 
out the eastern U.S. Young (1st-3rd instar) larvae of both species may frequently 
be found in the wild in Connecticut and Maryland in mid-September (L. a. 
astyanax commonly feeding on Prunus serotina Ehrh., and L. archippus on Salix 
spp. and Populus spp.). Twenty young archippus larvae were once collected at 
Middlefield (Middlesex Co.), Connecticut between 14-21 October 1968, all actively 
feeding prior to entering winter diapause (Platt, pers. obs. ). 

Developing larvae of both species are known to construct hibernacula (from the 
basal portions of tubular rolled leaves spun with silk) and to enter facultative diapause 
at third instar during the late summer and fall, in response to shortened daylength 
(Clark & Platt 1969, J. Insect Physiol. 15: 1951-1957). 

During the summer of 1966, three successive generations of L. arthemis-astyanax 
larvae were reared from a stock obtained from the intergrading population located 
at Shutesbury (Franklin Co.), Massachusetts (Platt & Brower 1968, Evolution 
22: 699-718). A total of 8 broods representing the three generations were bred and 
lab-reared under the ambient daylength for southern New England (Connecticut 
and Rhode Island) at room temperature. The incidence of diapause among the 
three successive generations of larvae increased dramatically from July through 
October (Table 1). 

Hong & Platt (in prep.) have determined that the critical photoperiod (that 
which induces 50% diapause among developing larvae) in Maryland and Vermont 
strains of L. archippus, lab-reared at room temperature, falls between 13.0 and 

VoLUME 28, NUMBER 1] 75 

TasLE 1. Incidence of facultative diapause among three successive generations of 
L. a. arthemis-astyanax butterflies from central Massachusetts—1966 data. 

No. of No. of Larvae 

Generation Broods Larvae Diapausing (%) 
1, early summer 
(July ) if 144 BAL 
2, late summer 
(August—early September ) 4 170 22,4 
3, Fall (October ) 3 66 98.5 

Note: All broods were lab-reared on Prunus serotina Ehrh. at room temperature under the 
ambient photoperiod for Connecticut and Rhode Island. 

13.5 hrs of light per 24 hr day. Photoperiods of 12 and 12.5 hrs per day induced 
diapause in 66-89% of the developing larvae representing these two strains. 

The ambient daylength at Durham, N. Carolina (approximate latitude = 36°N) 
decreases from 14:30 in mid-June to 13:25 in mid-August and down to 12:23 by 
mid-September (Duncombe 1966, The American Ephemeris and Nautical Almanac. 
U.S. Govt. Print. Off., Washington, D.C. 512 p.). On the basis of these data, and 
the diapause information given above, it is reasonable to assume that a high pro- 
portion of the larvae of both L. a. astyanax and L. archippus developing in the wild 
near Durham during August and September will enter diapause at third instar. Con- 
sequently, those larvae undergoing direct development in the late summer and 
fall will be relatively few. 

The matings which produced both of the North Carolina wild hybrids had to occur 
either in late August or (more likely) in September, a time of year when the adults 
of both species are at relatively low true population densities, despite their apparent 
local adult abundance noted above. The late capture dates of both hybrids makes the 
possibility of their successful backcrossing to parent-type females extremely remote 
(Platt & Greenfield, 1971). However, seasonably low late summer or fall temper- 
atures perhaps serve to make L. a. astyanax and L. archippus females more sluggish 
behaviorally, and hence less particular with regard to selecting their mates. In 
conclusion, a scarcity of adult butterflies of both species (i.e. mates of the same 
species ) may well contribute to a situation favoring natural interspecific hybridization 
in Limenitis. 

JosEPH C. GREENFIELD, Jr., Box 3246, Duke University Medical Center, Durham, 
North Carolina 27710. 

AusTIN P. Piatt, Department of Biological Sciences, University of Maryland Balti- 
more County, 5401 Wilkens Avenue, Catonsville, Maryland 21228. 


Recent descriptions of the widespread 1971 eruptions of the California tortoise 
shell, Nymphalis californica Boisduval, (Nymphalidae) (Powell 1972, Pan-Pac. 
Entomol. 48: 144; J. Lepid. Soc. 26: 226-228) have prompted this report of obser- 
vations in central California the following year. Weekly observations were made from 
4000 to 7200 feet along Interstate Highway 80 in the central Sierra Nevada in Nevada 
and Placer counties from 17 May—27 October and on 29 March; during intensive 
collecting in the Sacramento Valley (Yolo, Solano, Sacramento counties) at 10-100 
feet from 10 February—31 December; and frequently in the Vaca Mts., Yolo Co., from 
14 March-6 July and occasionally through 29 December. 


Scattered worn, hibernated N. californica were seen in the valley during the weeks 
of 6 and 13 March and in the Vacas on 14 March. No more were seen until the week 
of 8 May when large numbers of seemingly fresh butterflies appeared throughout the 
Vacas. While this flight was in progress, occasional, fresh single tortoise shells could 
be seen in the valley, mainly along its west edge near Vacaville and Winters. On 
26 May at 1700 hours a migration from west to east across the valley was observed 
from an elevated location just east of Davis, Yolo Co. Butterflies crossed a freeway 
overpass at the rate of one every 3 minutes, all headed due east. Most were flying 
from 30-60 feet above the vally floor, but a few were much lower. They continued 
moving through Davis from west to east most of the afternoon, at least until sunset. 
The movement was in progress again at 0900 the next day at comparable density. 
By evening it had dwindled somewhat, but stragglers continued to pass through 
Davis, moving from west to east, for about five days and a few individuals were seen 
at the east end of Yolo County, near the Sacramento River, as late as 27 June. On 
2 June only one tortoise shell was seen in the Vacas where there had been hundreds 
before the migration, and on 12 June none were seen at all. Thereafter no tortoise 
shells were observed in the Vacas in 1972 (although hibernators were again numerous 
in February and March of 1973). After 27 June none were seen in the valley until 

On 29 March numbers of hibernated tortoise shells were flying in the Sierran 
foothills and up to 3000 feet. On 17 May at Baxter, el. ca. 4000 ft., hundreds 
of fresh tortoise shells lined the roads and clustered about buildings. A few apparently 
hibernated ones were seen at the Marin-Sierra Camp, near 5000 ft., and at Castle 
Peak, 7200 ft. On 24 May an enormous migration was observed beginning at 0930 
hours, 4.5 mi. E Baxter (ca. 5400 ft.) and continuing for two miles of highway. 
The butterflies crossed the highway from S to N and moved generally upslope, 
passing at a rate of one every 5-10 seconds mostly from 3-10 feet above the ground. 
The migration did not reach to Castle Peak. Only a few live individuals were seen 
when we returned to the area at 1600 hours. Tortoise shells were abundant up to 
7200 feet on 31 May and 7 June, but at much lower densities than observed on 
24 May. At 5000 feet they disappeared entirely from 13 June-3 August. From 
3 August through 27 October they were continuously present again, but generally 
at low density. A movement occurred on 29 September when about 80 were observed 
moving downslope, E to W along I-80, flying into a strong headwind. These again 
appeared fresh. 

At 7200 ft. tortoise shells were seen every week from 17 May—4 October and 
again on 23 October. The first fresh specimens were observed on 30 June and 
thousands flew on 21 and 28 July. These seemed to be in local concentrations, mostly 
around towns, and no definite directional movement was noted. There were virtually 
none at this elevation on 3 August, when the species reappeared lower down. On 29 
September about 110 were seen at Boreal Ridge, mostly moving downslope and 
westerly along I-80, and a few were seen at Donner Pass. 

In the valley, N. californica was observed around Sacramento on 11 and 25 Sep- 
tember and 6 October. The first autumn sighting at Davis was on 3 October. There- 
after a few were seen each week through 23 October, and scattered sightings were 
made during the weeks of 13 and 20 November and 11 and 18 December. The 
bulk of records shifted from east Yolo Co. across the valley toward Vacaville and 
Dixon through the month of October. 

Tortoise shells began flying in the canyons of the Vacas the last week of January 
1973. Scattered worn individuals were noted throughout the valley from Winters 
to Sacramento beginning the second week of March. 

In summary, the seasonal changes in distribution of the California tortoise shell 
in central California in 1972 were: 

—A small flight of worn butterflies in the Vacas and valley and in the Sierra 
foothills in March. 



—A large flight of fresh butterflies in the Vacas in mid-May, followed by a mass 
movement from W to E across the valley floor in late May depopulating the Vacas 
for the rest of the season. 

—A large flight of fresh butterflies at 4000 feet in the Sierras in mid-May, develop- 
ing into a huge upslope migration in late May, disappearing in early June. 

—A large flight at 7000 feet in late July, disappearing in early August. Fresh 
butterflies, but no directional movement noted; concentrations local. 

—A small to moderate downslope migration at both 5000 and 7000 ft. on 29 
September, with scattered records of an autumn flight from late August through 
late October. 

—A small flight in the valley from mid-September into mid-December, mostly in 
October, with an east-to-west drift, presumably accounting for the spring 1973 
butterflies in the Vacas. 

Over-all, these records suggest that the California tortoise shell migrated eastward 
and upslope in spring, and westward and downslope in fall. If we do not postulate 
estivation or adult diapause, the generation sequence would be about as follows: 
a brood of new adults, progeny of hibernators, emerged in May in both the Vacas 
and Sierra foothills. The Vaca insects moved eastward across the valley floor (where 
N. californica does not breed, there being no Ceanothus) en masse in mid-May, 
while their Sierran counterparts moved upslope en masse at the same time. The two 
currents probably fused. The progeny of these insects emerged in July-August, with 
no well-defined migration, but perhaps a downward drift. The surge of butterflies 
moving downslope at the end of September may have represented a third, markedly 
smaller, generation. These would be the butterflies which appeared at low elevations 
at low densities and drifted across the valley floor in October. On this schedule 
each “brood” would take about eight weeks. (Generations of Milbert’s tortoise shell, 
N. milberti Latreille, take about seven weeks in New York.) 

During the 24 May eruption I examined 189 freshly killed specimens (103 
females). The females were all pre-reproductive, with no mature ova but substantial 
fat bodies. Of 35 examined for spermatophores, 32 were virgins. Only 11 females 
of the late July surge were collected, but all were also pre-reproductive. So, too, 
were 16 collected in the 29 September surge. On the other hand, most of the females 
collected between the big flights, when the species was at low density, were gravid 
and/or contained spermatophores, at least until early September. 

The brood sequence of N. californica is extremely obscure. In the complete absence 
of reports or observations of larval outbreaks in 1972 the number of generations 
can only be inferred from the condition of the adults. Although the data strongly 
suggest three broods, they do not rule out the possibility of only one—emerging in 
May, summering in the high Sierras, with only sporadic reproduction, and returning 
downslope to hibernate. Nymphaline adults are capable of extended periods of 
inactivity, but the reason for large-scale activity at eight-week intervals can only 
be described as arcane. Alternatively, one can postulate two “populations” in the 
Sierra, one resident and breeding without migration, one migratory and perhaps not 
breeding at all. 

Clearly, while the generalized picture Powell presents of a species erupting at 
irregular intervals from persistent epicenters (e.g. Mount Shasta) is broadly accurate, 
there were unexpected elements of regularity in the 1972 movements whatever the 
brood sequence that produced them. Following the 1971 outbreak N. californica appears 
to have colonized an extensive area and to have set up a migratory pattern which is 
seasonally adaptive. To judge by past history, its occupation of these areas will be 
temporary. The 1972 data raise some interesting questions: is it typical of mass 
movements that the females are not in reproductive condition? And how quickly do 
they come into condition? If females do not mate until after the migration, the 
chances of outbreeding are substantially increased—a possible genetic rationalization 
of eruptive periodicity (Brown 1957, Quart. Rev. Biol. 32: 247-279). 


Collections were made with the help of Dr. E. W. Jameson, Jr. and Mr. Allen 
Allison, both of the Department of Zoology, University of California, Davis, and 
Mrs. Adrienne R. Shapiro. 

ArtHuR M. SHapriro, Department of Zoology, University of California, Davis, 
California 95616. 


Between 0730 and 0810 MST, on an overcast day, 4 September 1970 (at 10 road mi. 
NE of Goblin Valley turnoff from Hwy. 24, on Hwy. 24, Green River Desert, Emery 
Co., Utah), Scott Ellis and I encountered Philotes rita Barnes & McDunnough spp. 
during a strong windstorm. We collected 19 rita clinging to stiff Ephedra plants in 
ca. 30-40 mph winds. The winds at first were calmer with no blowing sand but 
soon a gusty sandstorm from the SW hit with fine sand particles. Other plants in 
the area, including Eriogonum leptocladon Torr. & Gray var. leptocladon, rita’s 
foodplant here, were not nearly as upright in the wind as the Ephedra. Most of the 
rita perched in a head-down position (3—4 head-up) on the uppermost parts of the 
Ephedra, with the primaries tucked inside the secondaries, directed away from the 
wind’s angle, and buffeted by the wind. They clung on by their legs wrapped part- 
way around a stem. When approached or disturbed, some flopped down into the 
plant or onto the ground with the wind (they were still alert). These rita have been 
genitalically determined and represent an unnamed subspecies to be described 

OAKLEY SHIELDS, Department of Entomology, University of California, Davis, 
California 95616. 


Recent Letters 
Dear Dr. Sargent: 

In the last issue of the “Journal of the Lepidopterists’ Society” (Vol. 27, No. 3, 10 
August 1973, p. 210-219), there is an interesting study about the Biology of Prepona 
omphale octavia Fruhstorfer, by Alberto Muyshondt, of San Salvador, C.A. , presented 

“as this is the first time the life cycle of P. omphale octavia is fully described, . 
However, it is necessary to note that I published a long time ago (1933), sik the 
same species and surely the same subspecies, because the names of E. Le Moult 
(1932-33), in my opinion are subject to critical study and revision, in Guatemala 
occidental ( Department of Quezaltenango ). 

I am sorry I cannot send you a reprint of my work because I have only one in my 
library. The entire reference of this study is: Novitates Entomologicae. Paris. Fasc. 
3-4, 30 déc. 1933, p. 24-26, 1 pl. couleurs. “Observations biologiques sur les différents 
états de Prepona omphale guatemalensis Le Moult (Lép. Nymphalidae) par René 
Lichy.” Also in: Nov. Entom., janvier 1932, 2e. année No. 1, ler. supplément, p. 11, 
pl. couleurs. About the species, subspecies and aberrations described by E. Le Moult, 
cf. collection of “Nov. Entom.” 

I send this notice to you for the next “Journal.” Thank you very much. Very best 

Yours sincerely, 

RENE Licuy 


Chemin des Claies 

F-95320 Saint Leu-La Forét 


Dear Sirs, 

In a recent issue of your Journal (Vol. 27, no. 1, pp. 8-12, 1973) Professor Sargent 
concludes his short article with the following statement, “. . . numerous other experi- 
mental results (Sargent, 1968, 1969a and b) fail to support the reflectance matching 
mechanism proposed by Kettlewell (1955) and Ford (1964) to explain the selection 
of appropriate backgrounds by bark-like moths. On the contrary all of the evidence 
to date supports the view that these background selections are genetically fixed or 
innate responses.” 

I would like to ask one simple question: if in fact this statement is true, I would 
like Professor Sargent’s views as to how the two morphs of Biston betularia (£. typica 
and its melanic f. carbonaria) succeed so well in correct choice of backgrounds—two 
very different ones. 

Yours sincerely, 

Department of Zoology—Genetics Unit 
South Parks Road 

Oxford, England 

Ep. Nore: Since my views are solicited, I would suggest that the two morphs of 
Biston betularia differ both at the loci controlling the visible expression of melanism 
and at loci controlling background resting preference. I would assume that the 
different background preferences of the two morphs are fixed or innate, in the sense 
of being unmodifiable during the life of the insect. In such a situation, one would 
expect the evolution of mechanisms, such as the formation of supergene complexes, to 
insure that each morph inherits the appropriate background preference. 


Kenneth John Hayward was born on March 7th, 1891 in the small 
village of Pitney, near Taunton, in Somerset. At the age of eighteen he 
was already earning a living in London as an electrician, and by 1912 
he was working on the Aswan dam in Egypt in the same capacity. He 
joined the forces soon after the outbreak of war in 1914, serving in 
France, Greece and Cyprus, and returning to Aswan with the rank of 
Captain in 1919. In 1922 he returned to London and eventually secured 
a post as an engineer with the land-owning Argentine La Forestal 
company, which he took up in 1923 and held till 1929. It was during 
these years that he spent at Villa Ana and elsewhere in the Chaco that 
he amassed the very large collections that he presented almost in their 
entirety to the British Museum (Natural History), to be added to those 
he made whilst in Egypt. 

About 1930 he became associated with Albert and Adolph Breyer, 
both keen entomologists, working with them at Patquia, La Rioja, Argen- 
tina. His status as an entomologist, however, was only realized beyond 
doubt when in 1934 he was appointed in that capacity to the Agricultural 


Experiment Station of Concordia in Entre Rios, Argentina. From there 
in 1940 he transferred to the Agricultural Experiment Station at 
Tucuman, where his wanderings ceased. Here in 1944 he joined the 
Instituto-Fundacion Miguel Lillo of the National University of Tucuman, 
which conferred an honorary doctorate on him in 1950. Of recent years 
he enjoyed the title of Professor Emeritus. He died in Tucuman on 
May 21st, 1972. 

Hayward was a rather tall, spare man, somewhat reserved and seeming 
to be under tension from the sheer volume of work he always had in hand. 
At one time I used to receive from him with the greatest regularity, and 
much too frequently for my peace of mind, parcels of specimens and 
long numbered lists upon which I was required to fill in their names. 
I was unable to keep pace. Undeterred, he turned to others to supple- 
ment his identification service. It was only when, by these means, he had 
secured a firm basis that he began to make worthwhile contributions to 
our knowledge of the entomological fauna of the Argentine Republic. 

I don’t know enough about his publications to be able to evaluate them, 
but when I was Editor of the Entomologist (and ever since then) he 
regularly sent me notes on butterfly migrations in Argentina. W. H. 
Evans thought well of his work on Hesperiidae, considering the relatively 
limited facilities available to him. 

I am told that he was married, but “separated many years ago” and that 
he had a married daughter living in England with a son now at Uni- 
versity. An obituary was published by one of his associates, Dr. Willink, 
in Physis 81: 83 and another appeared in “La Gacete” S.M. de Tucuman, 
May 22nd. 

NorMAN D. Ritey, British Museum (Natural History), London, England. 


SP RT AOE peal eae ead 


Editor: THEoporE D. SARGENT, Department of Zoology, 
University of Massachusetts, Amherst, Massachusetts 01002 

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J. H. Masters, L. D. Mitter, A. P. Pratt, A. M. SHapimo, J. R. G. 


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Volume 28 1974 Number 2 


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Volume 28 1974 Number 2 


101 Avenida Norte #322, San Salvador, E] Salvador 

This is the third article in a series dealing with what my sons and I 
have found related to the life cycle and natural history of Rhopalocera 
encountered in the vicinity of San Salvador, capital city of El Salvador. 
As stated in our previous articles, the purpose of the series is to present 
the life cycle, including observations on the behavior of the early stages 
and adults, and to make known the foodplant of the local species of 
Neotropical Rhopalocera, as according to the literature many of them 
are incompletely known, and have been classified solely on the adult 
morphological characteristics. A major difficulty has been the identifica- 
tion of the species described. To overcome it, we have requested the 
help of Museums, mostly the Allyn Museum of Entomology, where Dr. 
L. D. Miller has made the identifications. 

This particular species has been placed through time in various genera 
by different authors: Cramer (1775) in Papilio, Hiibner (1807) in 
Consul, and Protogonius (1819), Duncan (1837) in Fabius, and Double- 
day (1844) in Helicodes. A host of specific names has been used too, 
among them the best known is hippona used by Fabricius (1777). We 
follow the name used by Comstock (1961), Anaea (Consul) fabius 
Cramer. Comstock leaves open the possibility that some subspecies 
might be valid. 

In our first article (Muyshondt 1973a) a summary description of the 
country and its climatic conditions was made. A. (C.) fabius is a dweller 
of wooded land, preferring ravines or creeks that cross coffee plantations 
(that are man made forests in this country). We have found the species 
from sea level to about 1500 m. In October 1969 we saw for the first time 
a female ovipositing on a plant that was identified as a Piperaceae, near 


Izalco, a town located about 45 km. W of San Salvador. In October 
1970, larvae in different stadia were found and collected near the village 
of Zaragoza (some 15 km. SSW of San Salvador), this time on two 
different species of the same Piperaceae. A few days later, in the same 
Zaragoza area, another female was observed ovipositing and eggs were 
collected, put in individual transparent plastic bags and brought back 
to our laboratory. Photographs were made of the eggs and the subse- 
quent stages of development until the adults emerged. Records of the 
developmental time, size and mortality were kept, and specimens of the 
different stadia were kept in alcohol. Since then the species has been 
reared during various months of the year to establish seasonal variations. 
Several species of Piperaceae, in addition to the ones on which the eggs 
and larvae were found, have been successfully used as foodplants. Breed- 
ing in all instances has been carried out under ambient lighting and 
temperature conditions. No moisture control was kept, but it was usually 
very high due to the fact that the material was kept in plastic bags, 
even though these were cleaned every day. 

Life Cycle Stages 

Egg. Translucent white with greenish tinge, almost spherical, with flattened base 
and depressed micropyle, surface smooth, and about 1 mm in diameter; all hatched 
IO eCaly Se 

First instar larva. Head light brown with darker markings, roundish, dispro- 
portionately large in relation to body, that is wedge-shaped from head to caudal end; 
grayish brown with scarce fine pilosity. After feeding on the leaf changes color to 
greenish-brown with tiny yellow markings. Measures upon emerging about 2 mm 
and grows to about 4.4 mm before moulting in five days. 

Second instar larva. Head black with tiny tubercles, white and yellowish, 
scattered mostly alongside lateral borders of epicrania. Two stubby horns on apex 
of epicrania. Body dark brown (almost black) with lighter peppering, thickening 
from first thoracic segment to first abdominal segment, tapering then gradually to 
10th abdominal segment. Grows to 0.9—1 cm in six days. 

Third instar larva. Head black with prominent tubercles, white or yellow, 
scattered in the area between ocelli and epicranial horns, which are now thicker, 
slightly longer and terminated in several short spines bearing thin setae. Body 
dark brown to black with heavy yellow sprinkling mostly on thoracic and mid- 
abdominal segments. On the caudal portion of the abdomen, sprinkling concentrated 
along spiracular area. Larvae grow to 1.5-1.8 cm in 4—5 days. 

Fourth instar larva. Head black with parallel vertical yellowish lines starting at 
adfrontal zone, the last and smallest located behind the ocelli. Head as thick as 
thickest abdominal segment. Body very dark brown or black with white sprinkling 
mostly concentrated on thoracic segments. Inconspicuous warts with tiny spines 
placed one at each side on subdorsal area of third thoracic segment. Scarce but 
noticeable setae alongside subspiracular area. White sprinkling on fourth abdominal 
segment and spiracular zone of caudal abdominal segments. Tubercles on head very 
abundant, most prominent of them yellow. Stubby horns with many tubercles and 
thick short spines with setae. Grows to 2.4—2.6 cm in 4—5 days. 

Fifth instar larva. Head black with some black and many yellow tubercles most 
prominent at sides of epicranium, and two near the upper adfrontal area. Frons 

VoLUME 28, NUMBER 2 83 

yellow, then yellow parallel vertical bands low on the epicranium. Horns as in 
fourth instar, but thicker and slightly longer. Body now dark green with dark red 
stains dorsally, mostly on thoracic segments and last abdominal segments. Spiracula 
yellow, surrounded by greenish ring. Spiracula on 2nd abdominal segment placed 
higher than the rest, same as spiracula on eighth abdominal segment. White pepper- 
ing scattered on body mostly on thoracic segments. Body now thicker than head, 
and short in relation to thickness. Grows to 3.5-3.8 cm in 9 or 11 days. 

Prepupa. Very thick and incurvated laterally, body all green. Duration two days. 

Pupa. Generally light green, occasionally light brown. Cremaster shining black. 
Abdominal segments taper sharply from wingcases (the thicker part of the pupa) 
to cremaster; thoracic segments taper gradually to slightly bifid head. Measures 
1.6-1.9 cm long, 1.2-1.3 cm laterally at widest point, and 1-1.1 cm dorsoventrally 
at thickest point. Duration 10-11 days. 

Adult. Shape of wings unique in genus. Forewing with projected angle starting 
at apex, going outwards to vein M 2, then sharply inwards to vein M 3, then slightly 
convex to tornus. Color dorsally dull black with conspicuous elongated yellow spot 
apically and row of three yellow elongated spots going from mid-costal area towards 
mid-outer margin, with an oval one under the last. Orange band covering basal 
and discal areas, parallel to black inner margin, not reaching tornus. Hindwing 
rounded with more or less spatulate tail on vein M 3, and sharp anal angle. Color 
orange except for dull black border alongside outer margin, with row of 4 yellow 
spots in black area, between tail and anal angle. The underside of both fore and 
hindwings is grayish-brown of varying shades, with no definite pattern. The body 
is orange, eyes reddish, antennae black basally, then orange turning to yellow, 
the tip usually black. 

No striking differences exist between the sexes, males somewhat smaller than 
females and having orange hairs alongside inner margin of hindwings. Much 
variation in shape of projected angle of forewing and of tails on hindwing, even 
among individuals emerged during same month. Total developmental time for this 
species varies from 45 to 50 days, females usually taking more time than males. 

Natural History 

All the plants on which we have found eggs and larvae of Anaea 
(Consul) fabius, and all the plants we have used as substitute food for 
the larvae, belong to the Piperaceae family. We have collected eggs 
and larvae on Piper tuberculatum, Jacquin, P. auritum H.B.K., and P. 
umbellatum L. and have used some others, not determined, with success 
when unable to obtain the original foodplant. Piperaceae in general are 
very common all over the country, among heavy second-growth plant 
communities, in humid ravines, along creek beds and coffee plantation 
roads. All the foodplants we have used have aromatic properties due 
to the content of essential oils, and usually have bitter flavor. 

The recently emerged larvae of A. (C.) fabius eat the egg shell com- 
pletely and stay under the leaf without further feeding for one day. 
Then the larvae move to the border of the leaf, usually to the tip, select 
a terminal of a vein, eat around it and prolong the vein with frass stuck 
with silk. The larvae use this as a perch when not feeding, and usually 
keep the head pointing outwards. This characteristic behavior is kept 
through the first, second and third instars. It sometimes happens that 


Figs. 1-9. Anaea (Consul) fabius Cramer: (1) egg, about 1 mm; (2) first 
instar larva recently hatched, about 2 mm; (3) first instar larva 4 days later, about 
4 mm; (4) second instar larva, about 1 cm; (5) third instar larva, about 1.7 cm; 
(6) fourth instar larva, about 2.4 cm; (7) fifth instar larva, two days after moulting, 
about 3.6 cm; (8) head of fifth instar larva; (9) prepupa, showing peculiar attitude. 


Figs. 10-14. Anaea (Consul) fabius Cramer: (10) side view of pupa, about 1.7 
cm long, 1 cm dorso-ventrally; (11) ventral view of pupa; (12) dorsal view of 
pupa; (13) adult, dorsal view, about 6.3 cm; (14) adult, ventral view. 

the whole leaf is eaten during this period, in which case the larvae move 
to another leaf where they make a new perch using the same system. 
During the fourth instar the larvae wander about the plant for two or 
three days, choose a larger leaf, and roll a portion of it, using silk, 
in the shape of a long funnel. From then on, until pupation, they remain 


inside of this funnel while not feeding, the head blocking the wide end 
and expelling the excreta through the narrow end. After feeding, usually 
done at dusk, the larvae come back to their hiding place, put the caudal 
end in position and crawl backwards into the funnel. When ready to 
pupate, the larvae abandon their funnel, wander about the shrub until 
they find a suitable place (a twig or a leaf, not always in the same plant, 
but always among heavy foliage), weave a small pad of silk, affix thereon 
the anal prolegs and stay there with the body incurvated laterally, not 
hanging. Just before doing this, the larvae expel an amount of greenish 
liquid mixed with excreta. The larvae of A. (C.) fabius through all 
instars are very slow moving, and when touched with a thin object make 
pushing movements with the tubercled head, and emit a pungent, though 
not disagreeable, scent, apparently from an eversible gland located 
anterior to the front thoracic legs. 

The pupae are either light green or light brown, regardless of environ- 
mental conditions, at least under laboratory conditions. We have 
simultaneously had green and brown pupae from larvae raised on similar 
diets, and among green leaves. The pupae are rather stiff and generally 
do not react when handled. At most, the pupae effect a short lateral 

The adults of A. (C.) fabius are, with the adults of A. (C.) electra West- 
wood, the slowest of all the Charaxinae found in El Salvador, even if, 
compared with other butterflies, they are rather fast. Males are very 
aggressive, and exhibit strong territorial defense behavior. They sit on 
top of a leaf, or at the side of a tree trunk, wings flapping from time to 
time, and will dash at anything flying near their resting place, whether 
it is another butterfly or just a falling leaf, then will return to the same 
or a nearby perch. Both sexes are very fond of feeding on fermenting 
fruits, sap of trees and even animal excrements. We have never seen 
them at flowers. 

When the females are ready to oviposit, they fly rapidly to the area 
where the foodplants are located, fly around one of the plants several 
times, and then approach the chosen one rather hesitantly, alight under 
a leaf, usually of medium development, and deposit one egg on the 
undersurface of it. They usually repeat the action on the same plant 
or on a neighboring one several times before flying away. 

Many times, when breeding this species, we have had tachinid larvae 
kill the larvae of the butterfly, generally when they reach the fifth 
instar or shortly after pupation. Some specimens of the adult of the 
tachinid have been sent to the U.S. Department of Agriculture for 
determination and these have been identified by Dr. C. W. Sabrosky 
as Chrysotachina sp. Another parasite found, even if very seldom, 


is a Chalcididae, determined by Dr. B. D. Burks, of the U.S. National 
Museum, as Spilochalcis sp., probably a new species. This latter parasite 
is polyembrionic and practically fills the pupa shell. In the case we sent 
for determination, 55 adults of the parasite emerged from one pupa. 
Apparently more than one egg had been injected, as males and females 
of the parasite were found. 

The larvae of A. (C.) fabius are very prone to a disease that softens 
their body until they burst and die. We have not witnessed any case 
of predation. 


According to Comstock (1961), the life cycle of Anaea (Consul) fabius 
has been at least partially described by several authors: Stoll (1787), 
Sepp (1852) (both under the name Papilio fabius), and Miller (1886) 
(under the name Protogonius drurii). Amazingly, Sepp mentions 
Mespilus americana (sic) as the foodplant. The genus Mespilus has 
been replaced, according to Standley (1922), by the genus Crataegus L. 
and belongs to the Malaceae (Apple family). In El Salvador at least, this 
species, Anaea (C.) fabius, feeds exclusively on Piperaceae. Was this a 
case of misidentification, or was a wandering pre-pupal larva or a pupa 
found in a nearby Mespilus? 

Apparently this is the first time a complete description of the life 
cycle of this species has been made, with photographs of the different 
stages as was the case with the descriptions of the life cycles of the 
other two Charaxinae, Prepona omphale octavia Frihstorfer and Anaea 
(Zaretis) itys Cramer (Muyshondt, 1973a, b). 

The egg of A. (C.) fabius is exactly like the egg of A. (C.) electra 
Westwood (with whom it shares the foodplant), Anaea (Memphis) 
eurypyle confusa Hall, A. (M.) pithyusa R. Felder and A. (Z.) itys 
(this last one is yellowish instead of greenish). The larvae are very much 
like the larvae of A. (C.) electra, even in coloration, and it is very hard to 
tell them apart until the fifth stadium, when the color of the head is 
lighter in A. (C.) electra. The larvae of A. (M.) e. confusa and A. (M.) 
pithyusa have the same shape as A. (C.) fabius but a completely different 
coloration. The larval behavior of the whole group of Anaea spp. 
mentioned, with the exception of A. (Z.) itys, is very similar in all 
instars from one to the other; they make the perch with the bared vein 
during the first stadium, and the funnel-shaped refuge during the fourth 
instar. During the pre-pupal stage all Anaea spp. we have reared behave 
alike: they do not hang like most Rhopalecera, but stay incurvated 
laterally, the body in contact with the supporting object. 


The species A. (C.) fabius exhibits a very effective defense mechanism 
based primarily on crypsis during the early stages: the translucent small 
egg is very hard to spot on the shadowy under-surface of the leaf; then 
the first, second and third instar larvae spend most of their time perched 
on the prolonged vein of a leaf, resembling to perfection a dried 
portion of it. The fourth and fifth stadia are spent hiding within the 
funnel-like contraption they make with their chosen leaf, and its entrance 
is blocked by their massive and tubercled head. The color and relative 
smallness of the pupae make it hard to locate among the profuse foliage 
of the shrubs. This cryptic behavior is common to most Anaea spp. found 
in this country, with the exception of A. (Z.) itys. But even this one, as 
well as the other Charaxinae we have studied, Prepona omphale octavia, 
behaves in the same manner up to the third instar. 

The adult of A. (C.) fabius, while in flight, can very easily be 
mistaken for a faster flying Licorea sp., Tithorea sp. or even an Heliconius 
telchinia Doubleday, all of which belong to families classically con- 
sidered unpalatable: Danaidae, Ithomiidae and Heliconiidae. This 
group is supposed to form a Miillerian mimicry complex. It is our 
opinion that A. (C.) fabius, feeding exclusively on Piperaceae, plants 
well-known for their content of essential oils and other at least bitter 
compounds, could very well have developed protective unpalatable 
characteristics, which augment its imitative coloration, and so effect its 
Millerian mimicry in this complex. This would explain the slowness of 
A. (C.) fabius in comparison with the other swift flying Anaea spp. 
But A. (C.) fabius adults do not solely rely on this defense mechanism: 
they also have the cryptic coloration of the underwings which makes 
the individuals inconspicuous among dry leaves. The species seems to 
enjoy a dual defense: unpalatability plus crypsis. 

It is to be noted that this duality of defense mechanisms seems to exist 
even during the larval stage. In addition to the cryptic behavior de- 
scribed above, the larvae, when molested, extrude a gland located an- 
terior to the prothoracic legs and emit a pungent scent. 

In spite of the complicated defense mechanisms of A. (C.) fabius, 
and the dusk and dawn feeding habits of the larvae, that minimize 
the risk of day-feeding predators, the mortality imposed on the species 
by ingestion-parasites is considerable. These ingestion-parasites are the 
Tachinidae that deposit their eggs on the leaf where the larvae are 
feeding. Regardless of the short developmental period (less than two 
months), which would allow no less than six generations a year, this 
species is rather scarce in the country, and mostly so during the rainy 
season. This fact leads us to deduce that parasites (Tachinidae in 
particular), are the principal factor that keeps the species in check. 


Parasites, being in general small animals, are, according to Janzen & 
Schoener (1968), much affected by dryness, such as is the case in E] 
Salvador from November to April. Thus it is during these months that 
A. (C.) fabius should be less affected by them and therefore should 
be more abundant. That is exactly what happens in fact. 


We are deeply grateful to Stephen R. Steinhouser for giving us access 
to his technical library and for sharing with us his own observations on 
adults of this species. To Dr. Lee D. Miller of The Allyn Museum of 
Entomology, who identified the species for us, and Dr. Theodore D. 
Sargent, who revised the manuscript, we express our gratitude. The 
eldest of this group is very thankful for the help and cooperation of his 
five boys, without which this study would not have been possible. We 
also thank Drs. B. D. Burks and C. W. Sabrosky for identification of the 
parasites mentioned. Specimens of early stages and adults have been 
deposited in The Allyn Museum of Entomology, Sarasota, Florida. 


Comstock, W. P. 1961. Butterflies of the American Tropics, the genus Anaea 
(Lepidoptera-Nymphalidae). Amer. Mus. Nat. Hist., New York. p. 51, 173. 

Cramer, P. 1775. Papillons éxotiques des trois parties du monde: L’Asie, 
L’ Afrique et L’Amérique. Amsterdam. Vol. 1. p. 141, 152. 

Dousiepay, E. 1844. List of Lepidopterous Insects in the Collection of the 
British Museum. London, Vol. 1. p. 112. 

Duncan, J. 1837. Foreign Butterflies. Nat. Library, Edinburgh. Vol. 18. p. 167. 

Fasricius, J.C. 1777. Genera insectorum, ete. Chilonii. p. 265. 

Htener, J. 1807. Sammlung exotischer Schmetterlinge. Augsburg. p. 148. 

1819. Verzeichniss bekannter Schmetterlinge. Augsburg. p. 100. 

Janzen, D. H. & T. W. ScHoreNnER. 1968. Differences in insect abundance and 
diversity between wetter and drier sites during a tropical dry season. Ecology 
49: 96-100. 

MUuuerR, W. 1886. Zool. Jahrb. Zeitschr. Syst. Geogr. Biologia der Tiere. Jena. 
Vol. 1. p. 503. 

Muysuonpt, A. 1973a. Notes on the life cycle and natural history of butterflies 
of El Salvador. I. Prepona omphale octavia (Nymphalidae). J. Lepid. Soc. 
27: 210-219. 

1973b. Notes on the life cycle and natural history of butterflies of El 
Salvador. II. Anaea (Zaretis) itys (Nymphalidae). J. Lepid. Soc. 27: 294-302. 

Sepp, J. 1852. Surinaamische Vlinders, Amsterdam. Vol. 3. p. 283. 

STANDLEY, P. C. 1922. Trees and Shrubs of Mexico (Fagaceae-Fabaceae). Cont. 
U.S. Nat. Herb., Vol. 23(2). 

Stott, C. 1787. Supplément, Papillons éxotiques des trois parties du monde. 
Amsterdam. p. 9. 



Department of Biology, Lawrence University, Appleton, Wisconsin 54911 

This paper summarizes a rearing study of Morpho peleides Kollar (Fig. 
1, as form limpida Butler) on the leaves of peanut, Arachis hypogaea L. 
(Leguminosae) under laboratory conditions in Costa Rica and Appleton, 
Wisconsin. It is generally known that the caterpillars of several South 
American species of Morpho feed on a variety of leguminous vines, 
shrubs, and trees (d’Aranjo e Silva et al., 1968). A recent study of the 
life history of Morpho peleides in Costa Rica and EI Salvador reports 
several papilionaceous legumes as foodplants of caterpillars (Young & 
Muyshondt, 1973). But there are no records of this butterfly feeding 
on peanut, which has a very widespread geographical distribution in 
the New World tropics (Leon, 1968). 

This study was undertaken primarily for the purpose of developing 
reliable and relatively easy methods for culturing the butterfly, as a 
prerequisite to experimental studies on the biochemical and behavioral 
aspects of feeding in Morpho caterpillars. The choice of Morpho peleides 
was made since mated females are very easy to obtain in the wild, and 
also because it is a member of the very frequently encountered achilles 
complex (or super species) in all of tropical America. 


The object of this study was to rear individuals from the egg through 
the adult stage. Eggs were obtained by confining a single healthy 
female butterfly in a 25 X 37 cm clear plastic bag containing a piece of 
fresh foodplant (usually Mucuna urens was used for this purpose). By 
repeating this procedure with several different females, a large number 
of viable eggs were obtained. Eggs were harvested from the leaves each 
day and the foodplant cuttings were replaced as they dried up. Females 
were removed from the bags and fed once or twice daily on juices from 
rotting banana. On the average, a mated female about 3-5 days old when 
caught lives three to four weeks in this manner and lays between 10 
and 105 eggs (an average of 65 eggs, N = 28 females) during this period. 
Eges were subsequently transferred to smaller plastic bags for hatching, 
keeping them at densities of usually 15 to 20 eggs. Eggs from each female 
were raised separately. All females used in this study as sources of eggs 
were wild caught at two mountain localities (1000 m. elev.) in the central 

VoLuME 28, NUMBER 2 91 


O cm 7 ey a & 

Fig. 1. Morpho peleides limpida Butler from Cuesta Angel, Heredia Province, 
Costa Rica: female (above), and male (below). About one-half natural size. 


TABLE 1. Developmental time (days) and some related ecological statistics for 
Morpho peleides on natural (Mucuna urens) and novel (Arachis hypogaea) food- 
plants in the laboratory.* 

Instar Instar Instar Instar  Instar 
2 3 4 5 

Statistic Egg Pupa Total 

Mucuna urens 
Devel. time (days)** 12.3 SO Ones A Bak yan 14.0 105.6 

Sake =O) EAN S212) E212) 40 eee 
Body length (mm) Del 1221.3) 2 6t ea ool ommmone 38.9 

Ske 0107) (S220 Fe 28) eG eee +24 
Head capsule 

width (mm) 2:0.) Dib 43) pio OS 

S.E. SEQ aaQp aa) sa(4t sell! 

N 275 Dil we DSi O43 OA Cems 240) 
No. which died 0 6 0 8 0 0 3 

Arachis hypogaea 

Devel. time (days) 12%3 13:05 oO: sell One ZO ote 14.4 104.8 

Sl Bic =O +22 +08 207 = 12) 20 0a 
Body lenpth (nm) 91 194 18A 986 380 73.0 sea 

Sea 210.0 == 013° 2016 == 0199-200) eles 
Head capsule 

width (mm) 2:0 2.55 AS ato eos 

Sa ssi seOJl se. se0.2) skO.S 

N 250 232) V232) 92215 Oia 220 
No. which died 18 0 TU 0 0 Il 0 

* The data are pooled here for measurements taken in San José, Costa Rica and Appleton, 
Wisconsin since results were very similar in both places. The raw data from each of these localities 
are, however, available upon request. The measurements of ecological statistics were always made 
on both foodplants simultaneously, so that all individuals were always exposed to the same 
environmental conditions (see text for a description of laboratory conditions). 

** See also Young & Muyshondt (1973) for other estimates of egg-adult developmental time and 
size range in Morpho peleides. 

highlands of Costa Rica (Cuesta Angel on the Caribbean slopes of the 
Central Cordillera, and Bajo la Hondura on the Pacific slopes). The 
butterfly is unusually abundant at both places, and females were easily 
baited with rotten fruit. 

A pilot rearing study was performed at Lawrence with a few eggs of 
M. peleides hyacinthus and M. polyphemus sent from El] Salvador by 
Alberto Muyshondt. These eggs were reared on peanut under green 
house conditions. The caterpillars, in second instar, were then trans- 
ported to Costa Rica and rearing continued on peanuts obtained locally. 
The success of this pilot study prompted the initiation of a larger scale 
rearing of peleides caterpillars simultaneously on Mucuna urens, a natural 
foodplant (control), and peanuts, a presumably novel foodplant for this 
species. This study was conducted in two parts: the first experiment 
was run in San José, Costa Rica, and the second one later in Appleton, 
Wisconsin (Lawrence University ). 

A total of 300 eggs was used for the Costa Rican study. These were 


Fig. 2. Thriving laboratory cultures of Morpho peleides and M. polyphemus 
on peanuts at Lawrence University: (A) M. peleides—fourth instar; (B) M. peleides 
—tifth instar; (C) M. polyphemus—fourth instar; and (D) an adult peanut ( Arachis 
hypogaea) plant (about 1% m tall) bearing two fourth instar Morpho caterpillars. 

obtained from 5 females, and all within a seven-day collection period. 
Each of 20 bags received 15 eggs. The bags were kept together on a large 
table away from direct sunlight, their positions on the table were changed 
frequently. Each bag received a code number. Room temperature was 
recorded daily during mid-morning. Foodplant was changed every four 
days and body lengths of caterpillars were measured usually every two 
days. Head capsules were always collected and stored separately for 


Fig. 3. Second-growth habitat of Morpho peleides at Cuesta Angel in Costa Rica 
(montane tropical forest): (A) high infestations of the caterpillars are frequently 
encountered on second-growth leguminous genera such as Mucuna and Machaerium 
which are very abundant along the sides of the road cut; (B) fifth-instar caterpillar 
in its cryptic resting position on a dead grass stem next to a Machaerium plant (16 


TABLE 2. A summary of some records for caterpillar foodplants in the genus 

Species Localities Genera Families Sources 
M. peleides Costa Rica Mucuna, Machaerium Leguminosae Young & Muyshondt, 1973 
Inga, Lonchocarpus 
El Salvador Machaerium, Inga Leguminosae Young & Muyshondt, 1973 
Trinidad Paragonia Bignoniaceae Barcant, 1971 
M. achilles Brazil Platymiscium Leguminosae d’Aranjo e Silva, 1968 
Machaerium, Dalbergia 
Pterocarpus, Myrocarpus Leguminosae Otero, 1971 
M. laertes Brazil Machaerium, Inga Leguminosae 
Luehea Tiliaceae d’Aranjo e Silva, 1968 
M. catenarius Brazil Acacia, Inga Leguminosae d’Aranjo e Silva, 1968 
Gymnanthes Euphorbiaceae d’Aranjo e Silva, 1968 
Scutia Rhamnaceae d’Aranjo e Silva, 1968 
Erythroxylum Erythroxylaceae d’Aranjoe Silva, 1968 
Cupania, Ratonia Sapindaceae d’Aranjo e Silva, 1968 
M. polyphemus El Salvador Paullina Sapindaceae Young & Muyshondt, 1972 
M. anaxibia Brazil Erythroxylum Erythroxylaceae d’Aranjoe Silva, 1968 
Nectandra Lauraceae d’Aranjo e Silva, 1968 
Clusia Guttiferae d’Aranjo e Silva, 1968 
Eugenia Mrytaceae d’Aranjo e Silva, 1968 
Ficus Moraceae d’Aranjo e Silva, 1968 
M. menalaus Brazil Erythroxylum Erythroxylaceae d’Aranjoe Silva, 1968 
M. hercules Brazil Abuta Menispermaceae d’Aranjo e Silva, 1968 
M. richardus Brazil Abuta Menispermaceae d’Aranjo e Silva, 1968 
M. aegae Brazil Bambusa, Chusquea Gramineae d’Aranjo e Silva, 1968 
M. portis _ Brazil Chusquea Gramineae d’Aranjo e Silva, 1968 

each bag. One half of the bags received Mucuna and the remaining ten 
received peanut. General day-to-day husbandry of the cultures also 
included removal of fecal material, dead caterpillars (recording the date 
of death), and periodic wiping of excess condensation. Three trained 
people performed the “sampling” of caterpillars and general husbandry, 
but the same person seldom sampled the same six or seven bags on two 
consecutive dates. Caterpillars were transferred as active prepupae 
to sturdy potted plants for pupation. Pupae were kept under the same 
room conditions as caterpillars and eclosion dates were recorded. Pupal 
size (length and width), but not weight, was recorded. Pupae of peanut- 
reared individuals were kept separate from those of Mucuna-reared indi- 
viduals. The wing-span of all emerging adults was also recorded. 

The same procedures were used for the subsequent study at Lawrence 
University, with the exception of a reduction in the number of cater- 
pillars studied. There were 125 caterpillars reared on Mucuna and 100 
caterpillars reared on peanut (seeds obtained from Olds Seed Co., Madi- 
son, Wisconsin). The cultures (Fig. 2) were kept in an air-conditioned 
laboratory whose mid-morning temperatures ranged from 21.8 to 24.0°C. 
The eggs used to establish the Lawrence cultures were obtained from four 

August 1972). Note: As of late March 1973, this section of road cut has been 
drastically widened by bulldozers, destroying a great deal of available roadside food- 
plants for Morpho. 


females captured in Costa Rica and brought to Appleton within a few 
days; the eggs were laid over an eight-day period. Prior to this time, 
thriving cultures of both peanuts and Mucuna (seeds brought from Costa 
Rica) were established at Lawrence for the sole purpose of rearing 
peleides and other Morpho. 


In the original pilot study, all of the caterpillars of peleides completed 
development successfully, but all of the polyphemus caterpillars died 
during the late fifth instar. Developmental time was not followed care- 
fully in this study. 

In the two major studies, there were no differences in the performance 
of caterpillars of M. peleides on Mucuna and peanuts (Table 1). Cater- 
pillars are apparently equally viable on both foodplants in the first 
generation. The various measurements given in Table 1 are adequate 
indicators of performance for caterpillars and pupae. The size range of 
adults reared on the two plants was very similar with no consistent trends 
towards increased (or decreased) wingspan on either plant. Very interest- 
ing is the similar success in rearing peleides in Costa Rica and Wis- 
consin (Table 1). Again, there were no consistent trends in the data 
supporting the view that rearing was more (or less) successful at either 
place. There was also no difference in the number of eggs in the bodies 
of virgin females reared on either foodplant: Mucuna-reared females 
less than two days old contained 61 + 7.5 (N = 55) eggs and peanut- 
reared females contained 60 + 5.8 (N = 46). Duration of older instars 
and the sizes of caterpillars and pupae were less variable for peanut- 
reared individuals (Table 1). 

Wild-caught healthy females in captivity will not lay eggs on peanut 
leaves while the same females will readily lay many eggs on Mucuna 
leaves under the same conditions. An attempt to obtain oviposition on 
peanut from peanut-reared mated females has not been done since I 
have been unable to achieve successful mating of peleides in captivity. 

In the pilot study on foodplant acceptance with Wisconsin legumes, it 
was found that second instar larvae readily accepted and survived on 
both Robinia and Gleditsia. This very interesting preliminary result 
will prompt me to conduct a large-scale controlled rearing study using 
several Wisconsin trees in the future. 

A representative portion of the known foodplants for the caterpillars of 
Central and South American (Brazilian) Morpho is given in Table 2. 
If we assume for the moment that Morpho and flowering plants evolved 

VoLUME 28, NUMBER 2 97 

at about the same time, the caterpillar-foodplant radiation of the genus 
can be discussed in a speculative but interesting manner. Based on 
present fragmentary knowledge of foodplants used by Morpho (Table 2), 
I propose that there were several different adaptive radiations within the 
genus, but that one of these was far greater than the others. Borrowing 
from the recent phylogenetic scheme of flowering plant evolution dis- 
cussed in Takhtajan (1969), the several “minor” foodplant radiations of 
Morpho included the families (in parentheses; see Table 2) in these 
orders: Ranunculales (Menispermaceae), Laurales (Lauraceae), Theales 
(Guttiferae), Urticales (Moraceae), Euphorbiales (Euphorbiaceae), 
and Poales (Gramineae ). But it is the derivative orders of the Saxifragales 
that formed the major basis for foodplant radiation in Morpho. The 
following orders and families very close to, or derived from, the Saxi- 
fragales (according to Takhtajan, 1969) contain foodplants of several 
Morpho (Table 2): Fabales (Leguminosae), Sapindales (Sapindaceae), 
Mrytales (Mrytaceae), Geraniales (Erythroxylaceae), and Rhamnales 
(Rhamnaceae). No other clear pattern of foodplant exploitation exists 
for Morpho since the minor groups are scattered across the phylogenetic 
scheme. Of course, this may be an artifact of the scheme proposed by 
Takhtajan; but departures would be minor and the same general pattern 
should result. Also note that the Rutales (which contains Rutaceae) are 
also derived from Saxifragales; in March 1973, I discovered several second 
instar larvae of peleides feeding on a vine in the Rutaceae in the under- 
story of a small semideciduous wet forest in Guanacaste, Costa Rica. 
Since the plant specimen was sterile, no further identification was made. 
These comments on foodplant radiation assume that the larval food- 
plant records of Morpho are accurate at the family level. It may be 
beneficial to re-check in the field some of the scattered records, espe- 
cially ones like Moraceae and Euphorbiaceae (Table 2). 

Thus it emerges that some species of Morpho, including members of the 
achielles complex (which includes peleides) not only feed on Legumi- 
nosae, but may in fact be preadapted to exploit other genera and species 
within this family. The data presented here for peleides on peanuts 
bear this out, if we assume that peanuts are not in fact used as foodplant 
in the wild. Such a preadaptation could result in species like M. peleides 
feeding on peanuts and other legumes. Being herbaceous, peanuts may, 
in fact, be an easier foodplant for digestion by caterpillars, as suggested 
by the reduction in the variability of developmental time and size during 
the ontogeny of M. peleides on this plant. This may be due to greater 
consistency of the leaves in this annual plant. In the wild, even very 
young caterpillars of M. peleides are found on very old and tougher 
leaves of foodplants (Young & Muyshondt, 1973). That M. peleides in 


particular may be especially preadapted for the exploitation of many 
different legumes is also suggested by the large number of foodplant 
species and genera that this species is found on locally in second-growth 
plant communities in Costa Rica. Here, the caterpillars are generally 
found on a variety of leguminous vines and small shrubs even along road 
sides where young second-growth is frequently encountered (Fig. 3). 
As with peanuts in the laboratory, the developmental time and size 
range of individuals reared on these different natural foodplants are very 
similar (Young & Muyshondt, 1973), indicating that the species performs 
equally well on all of these plants. If we assume that the deaths of the 
M. polyphemus caterpillars feeding on peanuts in the small pilot study 
was due to some metabolic or physiological incapacity to handle this 
food properly, it is possible that this species is less preadapted for 
generalized leguminous feeding than M. peleides. Partial support for this 
idea comes from the known foodplant data for M. polyphemus in El 
Salvador, and its close relative, M. catenarius, in Brazil (Table 2). The 
majority of foodplants are not legumes (Table 2) and these butterflies 
may have followed a different evolutionary path for foodplant exploita- 
tion from that of M. peleides and its close relatives. Of course, the data 
here for M. polyphemus are very preliminary and more extensive rearing 
tests on peanuts must be performed to demonstrate reduced performance 
on this plant. In light of these preliminary findings and their implications 
concerning evolutionary divergence in caterpillar foodplant exploitation, 
it could be very interesting to conduct similar rearing studies of other 
generally non-leguminous feeders of Brazilian Morpho (anaxibia, mena- 
laus, hercules, aega, etc—Table 2) with peanuts. 


(1) Caterpillars of the neotropical butterfly, Morpho peleides were 
reared in Costa Rica and Appleton, Wisconsin on Mucuna urens (a known 
natural foodplant) and peanuts, Arachis hypogaea, under identical con- 
ditions. While both plants are in the Leguminosae, the assumption was 
made that peanuts would be a novel foodplant for this butterfly since no 
records of it feeding on peanuts in tropical America are known. Further- 
more, all of the known leguminous foodplants of the butterfly are woody 
perennials and not herbaceous annuals. 

(2) Using various measures of performance such as egg-adult de- 
velopmental time and body size, it was found that caterpillars were 
equally viable on either plant. There was less variability in performance 
among caterpillars reared on peanuts. 

(3) A pilot study of rearing caterpillars of Morpho polyphemus on 
peanuts showed that they succumb in the fifth instar. But since a very 

VoLUME 28, NUMBER 2 99 

small number of caterpillars were studied, it could not be determined 
if these deaths were accidental or actually due to improper handling of 
the food by the digestive machinery of the caterpillars. This species in 
the wild feeds primarily on a variety of non-leguminous foodplants and 
it may eventually be shown that the caterpillars are less conducive to 
leguminous feeding. 

(4) Based on foodplant records and the phylogeny of flowering plants, 
it is speculated that the major adaptive radiation of Morpho occurred 
on plant families within various orders close to, or derived (in evo- 
lutionary time) from, the Saxifragales. Of these orders and families, the 
major array of foodplant exploitation is in the Leguminosae, a member 
of the Fabales. 

(5) The idea is advanced that M. peleides is preadapted to feed on 
many genera and species of legumes locally and there are some field data 
to support this view (Young & Muyshondt, 1973). Other legume-feeding 
species support this view (Young & Muyshondt, 1973). Other legume- 
feeding species of Morpho may show similar ecological flexibility while 
generally nonleguminous feeding species may not. 


This research was supported by a grant from the Bache Fund of the 
National Academy of Sciences (No. 120), and partially by National 
Science Foundation Grant GB-33060. Logistic support in Costa Rica 
was provided by the Costa Rican Field Studies Program of the Associ- 
ated Colleges of the Midwest (A.C.M.). Roger Kimber and John 
Thomason (Lawrence University) assisted with the rearing studies. 
Keith S. Brown, Jr., and Woodruff W. Benson read a revised version of 
the manuscript and made several helpful suggestions. 


Barcant, M. 1971. The Butterflies of Trinidad and Tobago. Collins, London. 

p ARANJO E Siva, A. G., C. R. Gonzatves, D. M. GatvAo, A. J. L. GONZALVEs, J. 
Gomes, M. NAscIMENTO Sitva & L. DE Stmont. 1968. Quarto catalogo dos 
insectos que vivem nas plantas do Brasil; seus parasitas e predadores. Ministério 
de Agricultura, Rio de Janeiro. 

Leon, J. 1968. Fundamentos Botanicos de los Cultivos Tropicales. Inst. Inter- 
Amer. Ciencias Agriculas de la OEA, San Jose, Costa Rica. 

Orrero, L. S. 1971. Instrucoes para criacao da borboleta “Capitao-do-mato” 
(Morpho achillaena) e outras especies do genero Morpho (“Azul-seda,” “Boia,” 
“Azulao-branco,’ “Praia-grande”). Inst. Brasileiro Desenvolv. Florestal (Rio 
de Janeiro ), 27 p. 

TaxurayaANn, A. 1969. Flowering Plants. Origin and Dispersal. Smithsonian Press 
(English translation ), Washington, D.C., 310 p. 

Younc, A. M. & A. Muysnonpr. 1972. Biology of Morpho polyphemus in El 
Salvador. J.N.Y. Entomol. Soc. 80: 18-42. 

. 1973. Notes on the biology of Morpho peleides in Central America. 

Carib. J. Sci. 13: 1-49. 



Joun H. MAstTers 
5211 Southern Avenue, South Gate, California 90280 

The most recent revision of the genus Speyeria (dos Passos & Grey, 
1947) and the most recent checklist for Nearctic butterflies (dos Passos, 
1964) have designated the southwest Manitoba population of Speyeria 
aphrodite (Fabricius) as subspecies mayae (Gunder). The name mayae, 
as proposed by Gunder, is unavailable and the name manitoba (Chermock 
& Chermock) must be used instead. 

Jean D. Gunder described mayae (1932) as Argynnis aphrodite cypris 
transitional form mayae from a pair of specimens collected by Marjorie 
May at Sand Ridge, Manitoba. Under the provisions of the International 
Code of Zoological Nomenclature (1961), this name is unavailable as a 
species group name because it was proposed as a quadrinomial and be- 
cause the author's intent was to describe an aberrant form and not a 
subspecies. Gunder coined the term “transitional form” to be used to 
refer to those types of aberrations that he considered nameable. Gunder’s 
holotype (Fig. 1, A & B) is a weird aberration of a sort that infrequently 
pops up in Speyeria. The Code provides, however, that although a name 
is unavailable when proposed, it can become available at a later date 
if elevated to a species group name. This was done when dos Passos & 
Grey (1947) elevated it to the subspecies rank. When a name is elevated 
in this manner it must take the date and the authorship of the elevation, 
in this case dos Passos & Grey 1947. 

In the meantime Chermock & Chermock (1940) described Speyeria 
aphrodite manitoba from the same locality: Sand Ridge, Manitoba. 
Their name has priority over mayae dos Passos & Grey. A typical looking 
male of Speyeria aphrodite manitoba is illustrated (Fig. 1, C & D). 

A very similar situation occurred with the southwest Manitoba prairie 
population of the Speyeria atlantis (Edwards) complex. Gunder (1927) 
described an aberrant as Argynnis lais tr. f. dennisi; this name being 
unavailable until being elevated by dos Passos & Grey (1947). Chermock 
& Chermock (1940) described Argynnis atlantis hollandi from nearby 
Riding Mountain, Manitoba. Their name, however, applies to the dark 
forest population of the Speyeria atlantis complex, and is not a subjective 
synonym of dennisi. It is my opinion that there are two species involved 
in what dos Passos and Grey call “Speyeria atlantis.” The name dennisi 
is available then, but must be credited to dos Passos & Grey 1947. 



‘azis JUNJoR sainsty [py “AWD YOR MON ‘Ar1OjSTET [eNJeNY FO UNosny UvoLoury sy} JO WOTaT[oo ey] UT o1e susUTTOedG “uoUTOeds 
aues JO episiopun (qq) ‘sruueq yor{ Aq poyeaT[oo ‘EEeBT “‘sNY FT “eqoyuRP, “YRneg ‘(yooursyD x» YooutteyD ) vgopupu ayposydo 
pisahadg ‘ayeut yeordAy (_)) ‘ueuttoads sures Jo spisiopun (gq) ‘Avy otofaeyy Aq peyejoo “TEE ‘3deg OT ‘oyxR'] wed sousg iveu “veqo} 
-TuryY espry pues, “1apuny apvfinw “Ff °31y siidho ayposydny swuhsuy ‘opeut adAjojoy (y) ‘:aupo1ydp piwahadg jo suoumloadsg 9=*T “SI 


In a discussion of these butterflies described from “Sand Ridge, 
Manitoba” some mention should be made as to the whereabouts of Sand 
Ridge; a locality that is not to be found on any map. Sand Ridge was a 
favored collecting locality of Jack May, Vern Harper and L. P. Baker 
and has become the type locality for a dozen taxa in Lepidoptera. The 
actual site is a gravel ridge, which was formed as a beach on glacial Lake 
Agassiz, 8 miles east of McCreary, Manitoba. Bener Dam Lake, a rather 
small impoundment is here. This locality is just east of Riding Mountain 
and is in western Manitoba. In the past many persons have placed “Sand 
Ridge” in the vicinity of the town of Sandilands or the Sandilands 
Provincial Forest Reserve. These are both in southeastern Manitoba on 
the other side of the Red River Valley/Lake Winnipeg divide that 
separates many species of Lepidoptera into eastern and western sub- 
species. It is very important then that anybody working with taxa de- 
scribed from Sand Ridge understand exactly where it is. 


I am grateful to Dr. F. H. Rindge of the American Museum for allowing 
me to examine specimens of the former Gunder collection, now a part of 
the American Museum collection in New York City. 


CuHERMOcK, F. H. & R. L. CHERMock. 1940. Some new diurnal Lepidoptera from 
the Riding Mountains and the Sand Ridge, Manitoba. Canad. Entomol. 72: 

“CopE’. 1961. International Code of Zoological Nomenclature adopted by the XV 
International Congress of Zoology. International Trust for Zoological Nomencla- 
ture, London. 176 p. 

GunpDER, J. D. 1932. New Rhopalocera (Lepidoptera). Canad. Entomol. 64: 
276-285. . 

pos Passos, C. F. 1964. A synonymic list of the Nearctic Rhopalocera. Mem. 
Lepid. Soc. 1. 145 p. 

& L. P. Grey. 1947. Systematic catalogue of Speyeria (Lepidoptera, 

Nymphalidae) with designations of types and fixations of type localities. Amer. 

Mus. Novit. 1370. 30 p. 

VoLUME 28, NUMBER 2 163 



Department of Biological Sciences, Stanford University, 
Stanford, California 94305 

An investigation of populations of Euphydryas editha Boisduval re- 
veals a disjunct distribution of foodplant choice (Fig. 1). Euphydryas 
editha may oviposit on plants of at least five genera: Collinsia, Castilleja, 
Pedicularis, Orthocarpus (Scrophulariaceae), and Plantago (Plantagina- 
ceae). With rare exceptions, only a single plant species is selected in 
each population, even though plants that are selected elsewhere may 
be abundant. This parallels observations of Downey & Fuller (1961) 
on Plebejus icarioides Boisduval. We have visited as many Euphydryas 
populations as possible, identifying primary hostplants of 50 by observing 
oviposition or by locating eggs or webs of prediapause larvae. Post- 
diapause larvae may move onto secondary foodplants and may even 
prefer these to primary hosts (Table 1). Oviposition preference in the 
laboratory is not necessarily the same as that in the field, and cannot 
always be used as positive evidence for placing a population in a par- 
ticular foodplant category. 

Our present knowledge of the distribution of hostplant choice (mostly 
in California) is summarised in Fig. 1. Though it is difficult to separate 
cause and effect, there are strong correlations between plant species 
chosen and a) timing of flight season, and b) type of community in- 
habited. Early-flying, coastal populations are Plantago-feeding, with 
some oviposition on Orthocarpus (EW, WS) and fewer on Collinsia 
(CS). Low altitude, late-flying populations in the chaparral belt of the 
Inner Coast Ranges are all on serpentine soils and utilise Pedicularis 
densiflora Benth. ex Hook. (plant identifications follow Munz & Keck, 
1959). Very close to a number of Pedicularis-feeding populations, but 
on a scree at 6900 feet in elevation in Mendocino County there is a single 
population (HM) feeding on a small and rather scarce annual, Collinsia 
greenei Gray. At similar elevations of the Sierra Nevada and in the San 
Bernardino Mountains we have records of small Collinsia species being 
utilised: C. childii Parry ex Gray at CP, C. callosa Parish at WK, and 
C. parviflora Dougl. ex Lindl. on both the east (SN) and the west (SL) 
slopes of the Sierra. It seems likely (S. O. Mattoon, pers. comm.) that 


TABLE 1. Primary and secondary foodplants of some E. editha populations. 

Oviposition plants Secondary Postdiapause 
in order of prediapause foodplants in order 
Population importance foodplants of importance 
JR P. erecta O. densiflorus P. erecta 
O. densiflorus 
EW O. densiflorus O. densiflorus 
P. erecta P. erecta 
LO P. insularis P. insularis 
P. hookeriana P. hookeriana* 
DP P. densiflora P. densiflora 
C. bartsiaefolia C. foliolosa 
C. affinis 
C. bartsiaefolia 
CP C. childii C. childii 
WK C. callosa C. callosa 
SN C. parviflora C. parviflora 
P. lanceolata. P. lanceolata 
SLi > C. parviflora C. parviflora 
P. semibarbata 
GH P. semibarbata P. semibarbata 
MC C. tinctoria C. tinctoria 
C. sparsiflora 
Lonicera interrupta 
Plectritis ciliosa* 
IF C. tinctoria C. tinctoria 
SAGs C. nana P. heterodoxus 
C. nana 
™™ C. nana C. nana 

* Order of importance may vary from year to year. 

populations of this type occur widely in Lassen and Shasta counties. 
In the southwestern Sierra these Collinsia-feeding populations are inter- 
spersed with colonies in the same general habitat (coniferous forest 
clearings; sandy, granitic soil) in which Pedicularis semibarbata Gray 
is utilised (GH, BM). Both P. semibarbata and C. parviflora are com- 
mon and well distributed, but E. editha populations seem to be few 
and widely scattered at these altitudes. Thus, as with P. densiflora 
in the Inner Coast Ranges, the distribution of E. editha is not limited 
by the distribution of its larval foodplants. 

At lower altitudes (1000-4000 ft.) in the western Sierra is a N-S belt 
of E. editha populations which are hostplant specific for Collinsia 

VoLUME 28, NUMBER 2 105 

tinctoria Hartw. ex Benth., rejecting even other Collinsia species where 
these are present (C. sparsiflora F. & M. at MC, and C. sp. at IF). 

Finally, at high altitudes (8000-11,500 ft.) along the crest of the Sierra 
we have found oviposition on Castilleja nana Eastw. to be the rule. 
At one of these populations (EP) we found another case of rejection of 
a congeneric plant, Castilleja breweri Fern. At EP, C. breweri is as 
abundant as C. nana and grows intermingled with it, but is not used 
for oviposition. 

Although the geographical range of E. editha extends from British 
Columbia to Baja California and eastwards to Colorado, Wyoming, and 
Alberta, we have little information on foodplant choice outside of Cali- 
fornia. In eastern Nevada we found two populations approximately 
three miles apart. In one of these, at 8000 ft. in a Pinon-juniper com- 
munity, Pedicularis centranthera is the foodplant, while in the other, 
at 11,000 ft., oviposition is on Castilleja lapidicola. In the McDonald 
Forest, near Corvallis, Oregon, we found postdiapause larvae feeding 
on the common weed, Plantago lanceolata L., in clearings of coniferous 
forest. We have been informed (D. V. McCorkle, pers. comm.) that 
this Eurasian import is also utilised for oviposition. 

We suspect that, even for California, the pattern of foodplant choice 
we describe here is incomplete. We have been unable to locate eggs 
or larvae of E. editha at a number of California populations where adults 
are well known, notably Parkfield Summit (Fresno-Monterey county 
line), Gold Lake (Sierra County), Bishop Creek (Inyo County), and 
Mather (Tuolumne County). Furthermore, several museum records of 
E. editha, such as those from eastern San Diego County, seem not to 
fit into any of the categories we have described. 

These data indicate that conclusions about foodplant relationships of 
an entire species of herbivorous insect should be made with caution 
when they are based on study of one or a few populations. Furthermore, 
since other aspects of the ecology of the insect, such as population 
dynamics, may be influenced by its choice of foodplant (White, 1973), 
these types of investigation also should ideally proceed on a population 
basis until a general pattern emerges. The lack (or complexity) of pat- 
tern in our data (Fig. 1) emphasizes the importance of evolution at 
the population level in the strategy of E. editha. Such evolution has 
allowed rapid exploitation of new food resources, such as the imported 
Plantago lanceolata and Plantago insularis Eastw. ( Bassett & Baum, 1969 ) 
as they have become available. This exploitive ability stems from the 
high reproductive potential of E. editha (Labine, 1968) and the low 
frequency of oviposition on alternative foodplants coupled with the 
ability to utilise these plants in response to selection. The hypothesis 


@ Plantago 
M@ Pedicularis 
O Collinsia 

DRAFT ress 2 
124 123 122 121 120 19 118 Ww 116 15 

Fig. 1. Euphydryas editha populations designated by their code initials. The 
symbols superimposed on this map of California represent the location and the 
larval foodplant of each population. 

that the range of plants acceptable to a migrant female E. editha 
broadens with increasing oviposition motivation as she searches (Singer, 
1971) would, if true, explain how a population can be founded on a 
foodplant which would not have been utilised in the parent population 
even if present. 

Since there is no simple correlation between primary hostplant used 
by an E. editha population and the subspecies to which the population 

VoLUME 28, NUMBER 2 107 

would be assigned, we propose to discuss the relationships between 
ecology and taxonomy of the insect in a separate paper. 


We would like to gratefully acknowledge aid, particularly in pinpointing 
E. editha population locations and flight times, from the following people: 
Ralph Wells, Fred Thorne, J. Tilden, William Swisher, Michael Smith, 
Oakley Shields, Harriet Reinhard, Paul Opler, James Mori, Andrew 
Moldenke, David McCorkle, Sterling Mattoon, Chris Henne, Lawrence 
Gilbert, Clifford Ferris, Thomas Emmel, Paul R. Ehrlich, Helen Cox, 
and David L. Bauer. This work was supported by NIH traineeships 
000-365-06 through 000-365-11; NSF grants GB 8038, 8174, 19686, 22853, 
and 35259; and by a grant from the Ford Foundation. 


Bassett, I. J. & B. R. BAum. 1969. Conspecificity of Plantago fastigiata of North 
America with P. ovata of the Old World. Can. J. Bot. 47: 1865-68. 

Downey, J. C. & W. C. Futter. 1961. Variation in Plebejus icarioides ( Lycaenidae ). 
I. Food plant specificity. J. Lepid. Soc. 15: 34-42. 

Lapine, P. A. 1968. The population biology of the butterfly, Euphydryas editha. 
VIII. Oviposition and its relation to patterns of oviposition in other butterflies. 
Evolution 22: 799-805. 

Munz, P. A. & D. D. Kecx. 1959. A California Flora. Univ. Calif. Press, Berkeley. 

Sincer, M. C. 1971. Evolution of food-plant preferences in the butterfly, Euphydryas 
editha. Evolution 25: 383-89. 

TuornE, F. 1970. Habitat: Euphydryas editha wrighti. J. Res. Lepid. 7: 167. 

WuirtE, R. R. 1973. Community relationships of the butterfly, Euphydryas editha. 
Ph.D. Thesis, Stanford University. 


THomas C. EMMEL 
Department of Zoology, University of Florida, Gainesville, Florida 32601 


Joun F. EMMEL 
1117 9th Street, Santa Monica, California 90403 

In the late 1960's, a new race of Papilio indra Reakirt was discovered 
in at least two isolated mountain ranges of the Great Basin state 
of Nevada. Described by Emmel & Emmel (1971), Papilio indra 
nevadensis is differentiated in the adult stage from all other known 










Fig. 1. Map of the known distribution of Papilio indra nevadensis Emmel & 
Emmel, including closely related segregates in Nevada, California, and Utah referred 
to in the text. The central area in Lander and Nye counties is the Toiyabe Range 
referred to in the text. @— closed circles = nevadensis populations in the Toiyabe 
Range and Toquima Range; © —open circle = Humboldt Range, Pershing Co., 
population; Mi — closed square = Westgard Pass, Inyo Co., California, population; 
[]— open square = Pine Valley Mountains, Washington Co., Utah, population; 
A, — open triangles = Grant Range and Quinn Canyon Range, Nye Co., popu- 
lations; A —closed triangles = Spring Mountains, Clark Co., population. 

VOLUME 28, NUMBER 2 109 

indra subspecies by the character combination of a wide postmedian 
yellow band on the forewing and hindwing, long tails, large size and 
elongated wings. The purpose of the present paper is to describe the 
distribution, habitat, behavior, foodplants, and life history of this dis- 
tinctive Nevada subspecies. 

Distribution, Habitat, and Habits 

Papilio indra nevadensis has been found in fair numbers in a series 
of canyons (especially Jett, Kingston, Peavine Creek, Summit, and Twin 
River Canyons) along the east side of the Toiyabe Range in Nye and 
Lander counties, Nevada. It also occurs to the east in the Toquima 
Range (Nye Co.) which runs parallel to the Toiyabe Range. Peter J. 
Herlan of the Nevada State Museum has taken two P. indra specimens 
in the Humboldt Range, Pershing Co., which represent either spring 
brood specimens of nevadensis or a population intermediate in adult 
characters between typical indra-and nevadensis. This mountain range 
is approximately 100 miles NNW of the Toiyabe Range. 

To the west of the Toiyabe Range, at Westgard Pass at the south 
end of the White Mountains in Inyo Co., California, a P. indra popu- 
lation is found which appears intermediate between nevadensis and 
typical indra. It utilizes the same Pteryxia petraea foodplant as nevaden- 
SESt oc! 

Southeast of the Toiyabe Range, several P. indra segregates which 
show a close affinity to nevadensis have been studied. P. indra larvae 
were collected on Lomatium parryi (Wats.) Macbr. (Umbelliferae) in 
the Grant Range and Quinn Canyon Range, Nye Co., in 1969, but the 
resulting pupae died, so the adult phenotype of these populations is 
not known. The coloration of these larvae appeared closest to that 
of P. i. martini. Farther south, in the Spring Mountains of Clark Co., 
adults and immatures of P. indra have been collected which show 
characters of both nevadensis and martini. The adults of this popu- 
lation are large with elongated wings as in nevadensis. The postmedian 
band of yellow spots tends to be intermediate in width between that 
of martini and nevadensis, and on the secondaries it tapers posteriorly 
as in martini. The color pattern of larvae from the Spring Mountains 
appears to be closest to that of martini. 

Another atypical P. indra segregate is found in the Pine Valley 
Mountains in extreme southwestern Utah. Adults and larvae of this 
population seem closest to those of the Spring Mountains’ populations. 
However, pupae from this locality are closest to those of P. i. kaibabensis. 

The locations of these populations are shown on the accompanying 
map (Fig. 1). 


Figs. 2-5. Habitat and foodplants of Papilio indra nevadensis: 2-3, Jett Canyon, 
from the east side of the Toiyabe Range, Nye County, Nevada; 4, canyon wall with 
scattered Pteryxia plants at Jett Canyon; 5, Pteryxia petraea at Kingston Canyon 
in the Toiyabe Range, Lander County, Nevada. 

The typical semi-arid, lower montane habitat where P. i. nevadensis 
occurs is exemplified by the Jett Canyon area in the Toiyabe Range. 
This canyon is located on the eastern slope near the southern end of 
the range (Fig. 2). The entrance to the canyon is very narrow, with 
steep walls on both sides of the narrow, 4-wheel-drive road going 
up the defile. The Canyon bottom is well watered by a permanent 
stream (Fig. 3). Typical vegetation within the canyon includes pinyon 
pine, willows (Salix exigua Nutt.), sagebrush (Artemisia tridentata 
Nutt.), Prunus virginiana L. var. demissa (Nutt.) Sarg., Purshia tri- 
dentata (Pursh) DC., and Holodiscus boursieri (Carr.) Rehd. The 
umbelliferous foodplants of this butterfly grow on the steep, rocky 
slopes and canyon walls (Figs. 4, 5). 

At present, we have only scant data regarding the spring brood. 
Based on our observations of immatures in June and July, we suspect 
that the spring brood flies in late May and June. The size of the sum- 
mer brood is variable; in 1967, 25 adults were collected in Jett Canyon 
in one day in August. In 1968, no adults were seen when the area 


was visited on 10 August, while on 3 August 1969, only one adult was 
taken. | 

Males were taken feeding on Cirsium species (thistles) and at mud 
or wet sand, and occasionally they visited blooms of Clematis vines. 
Females frequented Cirsium flowers and one was observed feeding 
on a Convolvulus (morning glory); several were flying along the canyon 
bottom. Other Papilio species actively flying at this time in these 
Toiyabe Range canyons are P. zelicaon Lucas, P. bairdii bairdii Edwards 
and P. b. form brucei Edwards, P. multicaudatus Kirby, and P. rutulus 
Lucas. Only the first species uses the same larval foodplant as P. i. 

The altitudinal span inhabited by P. i. nevadensis in the Toiyabe 
Range is 6200 to at least 7200 ft., with most specimens being taken 
between 6300 and 6800 ft. Undoubtedly hilltopping males ascend to 
the highest peaks of the Toiyabes, which are over 11,000 ft. 

Foodplant and Life History 

Throughout the Toiyabe Range and in the Toquima Range in central 
Nevada, Papilio indra nevadensis uses Pteryxia petraea (Jones) C. & 
R. (Umbelliferae) as a larval host. Females have been observed to 
oviposit on these plants in the field, and larvae of all five instars have 
been found on the Pteryxia in these mountain ranges. Pteryxia petraea 
is also found in the Humboldt Range, Pershing County, and doubtless 
serves as the foodplant for the P. indra population there. 

Egg: Globular in shape, smooth, about 1 mm, in diameter, and creamy white; 
laid singly on underside of foodplant leaf. Early instars black with white and 
light orange or yellow markings and closely resembling those of P. i. minori and 
P. i. kaibabensis. 

Fifth-instar Larva: Length: 40-45 mm. at maturity. Head: Width of head 
capsule, 4.0 mm. Head capsule pattern in most examples (Fig. 10) distinct from 
patterns of all other P. indra subspecies. Ground color black. Inverted “V” of 
light orange occurs on adfrontal margins, and low inverted “U” of similar color 
occurs laterally. On P. i. indra, minori, kaibabensis, and pergamus, these lateral 
marks extend dorsally to or near to top of head capsule. In P. i. martini and 
fordi, they are absent. 

Body (Figs. 6-9): Ground color black. In more common morph, each segment 
with narrow transverse cream colored band with pinkish tint, of a width covering 
two-thirds of anterior half of segment (not all the way to anterior edge) and 
ending on either side at level of spiracles (thoracic segments) or well below 
spiracles (abdominal segments). These bands yellowish to light pink in P. i. indra 
Reakirt (see Emmel & Emmel, 1973), white to pinkish gray in P. i. pergamus 
Hy. Edwards (see Comstock, 1928, and Emmel & Emmel, 1973), white, bluish 
white, or pale pink in P. i. fordi Comstock & Martin (see Comstock & Martin, 1955, 
and Emmel & Emmel, 1973), dull pink or salmon in P. i. martini Emmel & Emmel 
(see Emmel & Emmel, 1968), and rich bright pink in P. i. kaibabensis Bauer (see 
Emmel & Emmel, 1967) and P. i. minori Cross (see Emmel & Emmel, 1964). 
Transverse row of six rather large, yellowish orange spots located just beyond 


Figs. 6-10. Larvae of Papilio indra nevadensis: 6, common morph of fifth-instar 
larva, dorsal aspect; 7, common morph of fifth-instar larva, lateral aspect; 8, dark 
fifth-instar larva, dorsal aspect; 9, dark fifth-instar larva, lateral aspect; 10, head 
capsule pattern of fifth-instar larva, frontal view. 


posterior edge of each cream colored band in dorsal, suprastigmatal, and _ lateral 
positions. Thoracic legs and prolegs black; large white patch found laterally on each 
proleg and at base of each of the other segments. 

One very dark larva taken at Kingston Canyon (Figs. 8, 9). Here, cream bands 
and other patches quite whitish and reduced in size and yellow-orange spots on 
body very reduced (dorsal rows) or absent (suprastigamatal and lateral rows). 
Head capsule markings remain similar to those of lighter morph. 

Pupa: Length: 25-30 mm. Greatest width at wing cases: 7-9 mm. Morphologi- 
cally like those of other subspecies of P. indra. General ground color light dull tan, 
with mottling of various darker and lighter brown marks and lines over entire 


In the adult stage, Papilio indra nevadensis combines several of the 
key characteristics of P. i. pergamus (large size, elongated wings, long 
tails) and P. i. fordi (broad yellowish bands on the wings). In the 
original description (Emmel & Emmel, 1971), the new subspecies was 
said to be superficially closest to P. i. pergamus. Biologically, however, 
it differs in being double-brooded, having a generically different food- 
plant, and in having major larval and pupal color-pattern differences. 
There appear to be populations intermediate between P. i. indra and 
nevadensis to the north and west of the Toiyabes and intermediate 
between martini and nevadensis to the south. Populations in the Pine 
Valley Mountains in Utah show a combination of characters of nevaden- 
sis, martini, and kaibabensis. Thus the group of central Nevada popu- 
lations that are placed under the name nevadensis represent a true 
geographical subspecies which has departed evolutionarily in both larval, 
pupal, and adult characters, as well as general biology, from its con- 
specific relatives. 


The distribution, habitat, behavior, life history and foodplants of 
Papilio indra nevadensis are described from field work in Nevada, 
particularly in the Toiyabe Range. The mature larva differs in head 
capsule pattern, body pattern and coloration from larvae of all other 
P. indra subspecies. The foodplant is Pteryxia petraea (Jones) C. & R. 


The present paper is part of a continuing study of evolution in 
populations of the Papilio machaon complex in North America. We 
thank the Allyn Museum of Entomology and the Los Angeles County 
Museum of Natural History for travel funds from 1967 through 1969. 
Completion of this research was aided by National Science Foundation 
Grant GB-32151 as part of a study on chromosome evolution in 


Lepidoptera. Peter J. Herlan of the Nevada State Museum and his 
wife Barbara assisted greatly during field work in Nevada. Oakley 
Shields and Scott Ellis assisted in field collection of material. 


Comstock, J. A. 1928. Early stages of Papilio pergamus. Bull. So. Calif. Acad. 
Sci. 27: 82-86. 

& L. M. Martin. 1955. A new Papilio from California. Bull. So. Calif. 
Acad. Sci. 54: 142-150. 

EMMEL, J. F. & T. C. Emme. 1964. The life history of Papilio indra minori. J. 
Lepid. Soc. 18: 65-73. 

1966. A new Papilio from the Mojave Desert of California ( Lepidoptera: 

Papilionidae). Entomol. News 77: 57-63. 

1968. The population biology and life history of Papilio indra martini. 
J. Lepid. Soc. 22: 46-52. 

EMMEL, T. C. & J. F. Emmet. 1967. The biology of Papilio indra kaibabensis 
in the Grand Canyon. J. Lepid. Soc. 21: 41-48. 

. 1971. A new subspecies of Papilio indra from central Nevada (Lepidoptera: 

Papilionidae). Pan-Pacific Entomol. 47: 220-223. 

. 1973. The Butterflies of Southern California. Los Angeles Co. Mus. Nat. 

Hist., and Ward Ritchie Press, Los Angeles. 


Scott L. ELuis 
1217 41% St. N.W., Rochester, Minnesota 55901 

In recent years there has been vigorous interest in all aspects of the 
biology of North American species of Colias. Colias alexandra Edwards, 
a widespread species of the western foothills and mountains has attracted 
increased attention. Hovanitz (1950a) described its distribution, and 
(1950b) plotted frequencies of the dimorphic females. Ae (1959) in- 
duced laboratory crosses between C. alexandra and C. eurytheme Bois- 
duval. Masters (1970) and Ferris (1972, 1973) examined the taxonomy 
of the species. John M. Burns (unpubl.) has studied the electrophoretic 
variation of esterase in different Colias species, including alexandra. 
An attempt is made here to augment this work with notes on foodplants, 
population structure, and behavior in Colias alexandra. Observations 
were made on 35 C. alexandra populations during 1971 and 1972 in 
Colorado, Utah, Nevada, Idaho, and Montana. 

Colias alexandra is widely distributed along the axis of the Rocky 

VOLUME 28, NUMBER 2 115 

Mountains from New Mexico to Alberta and British Columbia, with 
outlying populations to the west in the Great Basin areas of Utah, 
Nevada, Idaho, eastern California, and then northward into eastern 
Oregon and Washington. East of the Rockies isolated populations occur 
in western Nebraska and the Black Hills of South Dakota. If christina 
Edwards is accepted as a subspecies of alexandra, the range is extended 
northeastward from Montana to Manitoba and northward to the Yukon 
River to approximately 67°N. For a distribution map and current 
taxonomic treatment, see Ferris (1973). C. alexandra is found in dry, 
open associations of the Transition and Canadian zones, most frequently 
from 7500-9000 ft. in the Colorado Rockies to 2000 ft. in northern Idaho. 

Oviposition Records. Edwards (1897) lists Thermopsis (Legumi- 
nosae) and Astragalus (Leguminosae) as natural foodplants. He found 
clover Trifolium repens L. (Leguminosae) to be a satisfactory laboratory 
host. Edwards received alexandra ova from several workers in Colorado. 
Edwards noted in his entomological journals that he received alexandra 
ova on 27 July 1884 laid on Astragalus from Nash at Rosita, Wet Mtns., 
Custer Co., Colorado. He also received eggs from Prof. G. H. French at 
Central City, Gilpin Co., laid 27 July 1886 on Thermopsis. Edwards (1873) 
reports that Mead observed alexandra ovipositing on Lupinus (Legumino- 
sae) in the northern part of South Park, probably in present-day Park Co., 
Colorado. From my experience with alexandra over much of its range, it 
appears that members of Lupinus are unlikely foodplants. Mead had only 
limited experience with the Colorado flora at the time of his observation, 
and may have misidentified the foodplant. McDunnough (1922) notes 
an oviposition by Colias christina in Alberta on a “small species of lupine 
with a greenish-white flower.” This vague description might apply to 
Astragalus canadensis L. var. mortonii (Nutt.) S. Watson, a foodplant 
for alexandra in northern Idaho and western Montana. Klots, in Ehrlich 
& Ehrlich (1961), lists Astragalus and Medicago as foodplants. Shields 
& Emmel (1969) observed oviposition on Astragalus miser Dougl. in the 
Wasatch Range in Sanpete Co., Utah. Ferris (1973) cites Astragalus 
serotinus as a foodplant. Barneby (1964) considers serotinus (Gray) a 
variety of Astragalus miser. 

Unpublished oviposition records unsubstantiated by herbarium de- 
terminations include Thermopsis pinetorum Greene from the White 
Mountains, Arizona, by Kilian Roever; Thermopsis divaricarpa A. Nels., 
Rampart Range Road, NE of Woodland Park, Teller Co., Colo., by F. M. 
Brown (photograph of ovum on plant). No ovipositions have yet been 
observed on Thermopsis montana Nutt., a species widespread west of 
the continental divide in Colorado. Kearney & Peebles (1951) consider 
T. pinetorum “doubtfully distinct” from T. montana. Mike Fisher ob- 


served oviposition on Oxytropis lambertii Pursh. (Leguminosae) near 
Parker, Douglas Co., Colorado. 

Foodplant-butterfly Relationships. Repeated ovipositions on a 
plant species in one locality, or oviposition on the same plant species 
in different localities is considered here to be a strong confirmation that 
the oviposition plant is the foodplant. No obvious oviposition mistakes 
were seen. Astragalus eremiticus is the only foodplant listed for which 
there is only a single oviposition sighting. 

Many members of the genus Astragalus and Oxytropis are known to 
accumulate toxic compounds in their systems, especially the element 
selenium. Colias alexandra oviposited on Astragalus miser, A. bisulcatus, 
and Oxytropis lambertii, which are frequently poisonous to livestock. 
The toxicity of A. miser appears to vary as a function of the selenium 
concentration of the many different types of soil the plant inhabits. 
Barneby (1964) notes that “miser var. oblongifolius is often browsed, 
even where innocuous feed is plentiful, at least in the Colorado Rocky 
Mountains.” For A. bisulcatus, Barneby (1964) states that: “On warm 
days and while drying in the press, the herbage gives out a strong smell 
of selenium disagreeable to most people and to some actually nauseating. 
It is one of the most dangerous and widely dispersed of the seleniferous 
stock poisons.” A study of the biochemical relationship between C. 
alexandra and A. bisulcatus might prove rewarding, particularly if 
selenium is metabolized and stored in the immature and adult stages of 
the butterfly. 

The ranges of C. alexandra foodplants often greatly exceed the range 
of the butterfly. Of particular interest is the absence of alexandra in 
most parts of Arizona, especially on the Kaibab Plateau north of the 
Grand Canyon (Kilian Roever, pers. comm.) where varieties of A. 
lentiginosus, A. miser, and A. bisulcatus are known to occur. C. alexandra 
is known from the nearby mountains in southwestern Utah. Astragalus 
canadensis has one of the widest ranges of any North American As- 
tragalus, embracing nearly the entire eastern half of the United States, 
ranging south to the coast of Texas. A. lentiginosus is a highly poly- 
morphic species that lives in a great range of environments and altitudes, 
and includes several varieties that occur in the deserts of southern 
Arizona and California. It is possible that C. alexandra has a physiologi- 
cal intolerance to the higher temperatures of desert areas, although a 
population at 5600 ft. in the arid Henry Mountains of southern Utah 
indicates that alexandra should be sought in comparable environments 
in northern Arizona. Much of eastern Colorado, Wyoming, South Dakota, 
eastern Montana, and Alberta remain for further foodplant investigations. 

Many members of the genus Astragalus are pioneer or “fugitive” spe- 


cies, often colonizing barren ground unacceptable to other plant species, 
and then disappearing as more competitive species enter the area 
(Barneby, 1964). Several parallel cases of local abundance of a C. 
alexandra foodplant appeared to be the result of disturbance of the plant 
community by man and his animals. In the La Sal Mountains of Montrose 
Co., Colorado, large stands of Lathyrus leucanthus were found growing 
next to stumps of Pinus ponderosa Lawson cut within the previous two 
years. In Piute and Juab Counties, Utah, Astragalus lentiginosus was 
found most abundantly along road cuts and in areas where sage 
Artemesia tridentata had been cleared. C. alexandra females were ob- 
served ovipositing on A. lentiginosus growing in the rubble of an aban- 
doned highway. A. miser at Black Canyon, Montrose Co., Colorado, was 
found in eroded gullies and in sage flats heavily grazed by cattle. 

It seems reasonable to assume that Colias alexandra must be a highly 
mobile species in some parts of its range as it follows the expansion and 
contraction of populations of its. Astragalus foodplant. Evidence from 
capture-recapture studies at Gothic, Colorado, where alexandra feeds 
on L. leucanthus, indicates that alexandra has a strong tendency to 
disperse as compared to the more sedentary Colias meadii Edwards 
(Ward B. Watt, pers. comm.). It was found from extensive travel over 
the range of alexandra that this species generally occurred in widely 
dispersed populations, with occasional local, large concentrations. A 
“large concentration” was one in which the density of adult alexandra 
was estimated at 20 or more individuals/100 m?. A large concentration 
of C. alexandra was nearly always accompanied by a local abundance 
of a foodplant. A “local abundance” was one in which patches of the 
foodplant occurred at high densities within small areas that ranged from 
100 m? to several km*. Astragalus canadensis in Idaho was found in 
densities up to 100 stems/m” in some patches. In dry areas of Utah, 
clumps of A. lentiginosus were scattered, but individual plants were 
often 0.2 m or more wide. 

Timber management practices in northen Idaho and Montana offer 
an explanation for the changes in population size of Astragalus canaden- 
sis, and for the spotty occurrence of C. alexandra. Closely related to the 
Siberian A. uliginosus L., A. canadensis var. mortonii is a conspicuous 
species, often reaching 0.8 m in height, with a head of greenish-white 
flowers which later form a cluster of small, erect ellipsoid pods. The 
outstanding characteristic of canadensis is its occurrence in large patches 
as a result of vegetative spread by rhizomes. Patches of the species 
are easily seen while driving at high speeds along the highway. A. 
canadensis grows most abundantly in lodgepole pine, Pinus contorta 
Dougl., forest that has just been logged and cleared. Dense stands of 



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canadensis were observed growing in the ashes of burn piles, and in 
still-visible tractor tracks. Young stands of P. contorta are thinned 
for optimum growth, and A. canadensis and many other species of herbs 
cover openings in the thinned forest. A. canadensis becomes much less 
frequent in mature or stagnant lodgepole pine stands, or in dense Douglas 
fir, Pseudotsuga menziesii (Mirb.) Franco—Western red cedar, Thuja 
plicata Bonn, forest. Direct comparisons for butterfly abundance were 
made between adjacent logged and unlogged stands. C. alexandra was 
found commonly only where A. canadensis was abundant, although large 
patches of A. canadensis were found where no C. alexandra were seen. 
It seems possible that present timber maintenance methods have re- 
placed a natural fire cycle which opened up clearings for invasion of 
A. canadensis, an early seral stage plant. 

The single factor of foodplant abundance as a cause of large alexandra 
populations is insufficient in itself. Increased nectar sources, and an 
open habitat for increased adult interaction may contribute to large 
populations. It was noted that Lycaena mariposa Reakirt, Lycaeides 
argyrognomen Bergstrasser, and Colias interior Scudder were often ex- 
tremely common in northern Idaho forest openings where C. alexandra 
was abundant. 

W. H. Edwards discovered that C. alexandra enters a 3rd instar 
diapause. This may be an adaptation in response to desiccation of the 
foodplant. In Juab and Piute Counties, Utah, at the end of June 1972, 
Astragalus lentiginosus var. araneosus was found in an advanced stage 
of desiccation, with most small plants completely withered. A few large 
plants, on which C. alexandra was ovipositing, held mature pods on some 
stems, and were still blooming on others. No new growth was observed, 
and leaves were hard and leathery. With high evaporation rates and 
scant rainfall in this part of Utah, the risk of drought is great during 
the summer months. Even in an area of comparatively high rainfall 
such as Gothic, Gunnison County, Colorado, C. L. Remington (pers. 
comm.) has noted that Lathyrus leucanthus withers toward the end 
of July. 

There was little opportunity for field observations on choice of food- 
plants by alexandra. In most populations where ovipositions were ob- 
served, only one species of Astragalus or Lathyrus was found, although 
search for other species was made. In many areas two or more accept- 
able foodplants surely must occur within the range of one alexandra 
population. In the Schell Creek Range of White Pine Co., Nevada, 
Barneby (1964) records varieties of Astragalus lentiginosus and A. miser 
within the area where alexandra was found ovipositing on A. eremiticus. 
In Cassia County, Idaho, at 5000 ft. on 3 July 1972, C. alexandra was 

VoLUME 28, NUMBER 2 121 

found abundantly in a small field (75 X 75 m) of alfalfa, Medicago 
sativa L. Between 1000 and 1200 MST alexandra was estimated at a 
density of 150-200 individuals in the field at one time. Such large 
numbers of this species had not been encountered previously, and it 
occurred to me that alexandra might be using Medicago as a foodplant 
in this locality. Careful observation of nearly thirty females during the 
prime oviposition time of late morning revealed only nectar-seeking 
behavior, and no oviposition or courtship behavior. Scattered Astragalus 
lentiginosus var. salinus plants were found along the road cut, and among 
sage shrubs bordering extensive sage and juniper flats. This discovery 
led me to believe that the source of the alexandra population was a 
colony of the Astragalus growing in the sage flats, with alexandra adults 
moving into the alfalfa field to feed. 

Voltinism. Colias alexandra is a univoltine species over most of its 
range. Emergence times vary for different populations, with western 
Colorado ones generally appearing several weeks before those in the 
Great Basin, Arizona, and in the northern Rockies. Lengths of emergence 
vary from 3 weeks in populations from dry areas in western Colorado 
to 2 months along the Front Range in eastern Colorado. 

Mike Fisher discovered a bivoltine alexandra population on the high 
plains near Parker, Douglas Co., Colorado. Confining dates for the two 
broods are, first brood: 25 May-—22 June; second brood: 25 July—19 
August, with no stragglers between broods. C. alexandra oviposited on 
Oxytropis lambertii in this area, although Astragalus species may be 
used as well. 

Interspecific Relationships with other Colias. Throughout its range 
C. alexandra is almost always sympatric with at least one other species 
of Colias, and in some areas up to four. C. alexandra and C. philodice 
are most often sympatric, although the peaks of their broods are not 
always synchronous. Ae (1959) states that in Colorado opportunities 
for interspecific mating are rare between widespread Colias species 
(philodice and eurytheme) and the “northern” species (alexandra, scud- 
deri Reakirt, and meadii) due to seasonal isolation. In western Colorado, 
the peak of alexandra’s flight period occurs between the spring and 
summer broods of C. philodice. On Mesa Verde, Montezuma Co., on 
26 May 1972, the alexandra:philodice ratio was 5:1, and on 16 July 
1971 in the same locality, the same ratio was 1:8. In Cassia Co., Idaho, 
in an alfalfa field on 3 July 1972, the alexandra:philodice ratio was in 
the range of 100:1, with no C. eurytheme seen. In western Colorado 
C. eurytheme is quite rare when C. alexandra flies in late June, but 
becomes more common after alexandra disappears. Throughout the 
Great Basin alexandra flies with both C. eurytheme and philodice, and 


in northern Idaho is sympatric with C. interior as well. Near Lake Pend 
Oreille, Kootenai Co., Idaho in thinned lodgepole pine forest on 14 
July 1972, Colias collections were made along a 0.2 mile strip of logging 
road from 0900 to 1200 PST. In this limited area nearly 200 C. alexandra 
were seen and collected, as well as 10 C. interior, 6 C. eurytheme, and 
2 C. philodice (all specimens seen of the last 3 species were collected). 
Several unsuccessful attempts by alexandra males to copulate with in- 
terior females were observed in this locality. Where alexandra and 
interior are sympatric, alexandra frequents open areas along roadways 
and meadows, while interior is found more often in the shade of pine 
woods. C. interior invades meadow habitats in localities where interior 
is more common than alexandra. 

In Canyon Creek Canyon, Ochoco Mountains, Crook Co., Oregon, 
where C. alexandra and C. occidentalis Scudder are sympatric, A. O. 
Shields (in litt.) notes that the two species are easily separable on the 
wing, and that there are other behavioral differences. C. occidentalis 
“was more confined to openings and edges of woods than alexandra, 
though both taken commonly in open, broad meadows.” In Canyon 
Creek Canyon on 10 July 1970, Shields observed oviposition by C. oc- 
cidentalis on the leaf underside of Lathyrus lanszwertii Kell. (Shields 
#109, det. J. T. Howell, Calif. Acad. Sci.). Shields saw a species of 
lupine Lupinus latifolius J. G. Agardh. (Shields #86) in Canyon Creek 
Canyon, the same foodplant species used by C. occidentalis at Camp Ellen- 
dale, Glenn County, California. 

Adult Nectar Sources. A study of nectar sources for several popu- 
lations of C. alexandra indicates that the insect visits a variety of plant 
species. Throughout western Colorado and parts of Utah the greatest 
concentrations of alexandra were found on various species of thistle 
Cirsium (Compositae). One large population on the South Rim of 
the Black Canyon, Montrose Co., Colorado, was found almost exclusively 
on Canadian thistle, Cirsium arvense L. This plant population had 
been introduced during the previous five years after the construction 
of a new road. On 29 June 1971, on the Uncompahgre Plateau, Montrose 
Co., Colorado, alexandra was observed to ignore Cirsium sp. in favor of 
mules ears, Wyethia arizonica Gray (Compositae). In Cassia Co., Idaho, 
alexandra chose the blossoms of Medicago sativa and bindweed Con- 
volvulus arvensis L. (Convolvulaceae) over those of a Chrysothamnus 
species (Compositae) which attracted Satyrium fuliginosum Edwards, 
Speyeria zerene Boisduval, Cercyonis oetus Boisduval, and Hesperia 
harpalus Edwards. In northern Idaho, Cirsium sp. was completely 
ignored, and alexandra was found most commonly on its foodplant, 
Astragalus canadensis var. mortonii. 

VOLUME 28, NUMBER 2 123 

Nectar sources are especially significant in concentrating alexandra 
populations in arid country. The widespread introduction of weeds, 
especially Cirsium sp., along roadcuts may have local effects on the 
density of alexandra populations. At Jericho Turnoff from Hwy. 6-50, 
5400 ft., Juab Co., Utah on 25 June 1972, C. alexandra was collected 
on two small thistle patches (tentatively identified as Cirsium vulgare 
L.) along an abandoned highway. This locality was extremely arid, with 
extensive sage flats changing to sand dunes a few miles to the west. 
No moisture in the form of mud or streams was available. The only 
other nectar sources utilized in this area were a few scattered blossoms 
in Astragalus lentiginosus clumps, and the flowers of a small introduced 
mint growing on the roadcut. It appeared that the thistle patches were 
recent introductions, owing to the lack of previous years’ stalks, and 
the absence of other thistles for many miles in all directions. Twenty-five 
C. alexandra were collected on thistle flowers from the two patches. A 
return to the area the next day netted only four specimens. Subsequent 
travel north, south, and west for several miles in each direction indicated 
that alexandra was either very scarce, or non-existent beyond 0.2 mi. 
from the thistle patches, although scattered A. lentiginosus clumps 
were seen along the Little Sahara Sand Dunes road to the west of 
Jericho. It appeared that we had collected nearly the entire emerged 
population from a large area, indicating that the thistle blossoms were 
a powerful attractant. This observation raises the question of whether 
adult nectar sources may be a limiting factor in the size of arid-land 
alexandra populations. In some areas, alexandra may be limited to the 
flowers of its foodplant, a situation shared by some desert-dwelling 
Philotes, and Apodemia mormo Felder & Felder, which feed on fall 
blooming Eriogonum species (Polygonaceae). 

Behavior. Colias alexandra followed a consistent behavioral pattern 
over its range. In open country on warm days individuals of both sexes 
arrived at nectar sources about 1030. Males were least wary at this 
time, and most easily collected. After 1200, both sexes began to leave 
the nectar sources. Males congregated on mud, or flew continuously 
over meadows or along the edge of the forest. Oviposition by females 
occurred most often between 1000 and 1330. The fast flight and large 
size of alexandra made it easy to separate from other Colias species 
on the wing. Beak-marked individuals were very rare, and no attacks 
by avian predators were observed. Although hundreds of alexandra 
were observed on nectar sources, no copulating pairs were found. This 
suggests that courtship and mating may take place at a distance from 
nectar sources. C. alexandra moved away from open areas during the 
hottest hours of the afternoon, and then another brief nectar feeding 


period occurred in some populations at 1600., On the Uncompahgre 
Plateau in San Miguel Co., Colorado, an individual was flushed from 
inside a sagebrush clump at 0800 MST where it had apparently spent 
the night. 


1. Colias alexandra oviposits on members of at least four genera of 
the Leguminosae: Thermopsis, Astragalus, Oxytropis, and Lathyrus. 
Lupinus is considered a very doubtful foodplant. Clover, Trifolium 
repens, is known to be a laboratory host. Field observations indicate 
that alfalfa, Medicago sativa, is an unlikely foodplant. 

2. C. alexandra oviposited on two species of Astragalus, and one of 
Oxytropis which are known to be toxic to livestock. At least one species, 
Astragalus bisulcatus, is known to be a selenium accumulator. 

3. C. alexandra foodplants are characteristically perennial, and show 
a scattered or patchy distribution. Several are dependent on the ayvail- 
ability of disturbed plant communities and early seral stages in forests 
for their optimum growth. Two species, Thermopsis divaricarpa and 
Astragalus canadensis, form dense patches due to spread by rhizomes. 

4. C. alexandra occurs most frequently in widely dispersed popula- 
tions. Occasional large, local concentrations are found. It is suggested 
that these large, local alexandra populations are primarily dependent on 
the local abundance of a foodplant. Other factors such as an open 
habitat and increased nectar sources may contribute to the support of 
large alexandra populations. 

5. Although univoltine over most of its range, a bivoltine alexandra 
population is known from the High Plains-Front Range contact area 
in Douglas Co., Colorado. 

6. C. alexandra is nearly always sympatric with one or more species 
of Colias. Limited data suggest that there are temporal and behavioral 
differences between alexandra and other species of Colias. 

7. Available nectar sources, particularly introduced weeds, may 
strongly influence density and size of alexandra populations occurring 
in very arid areas. 


The author is grateful to Arthur C. Allyn and Lee D. Miller of the 
Allyn Museum of Entomology, Sarasota, Florida, and to John M. Burns 
of the Museum of Comparative Zoology, Harvard University, whose 
support made travel and research possible during the summers of 1971 
and 1972. F. M. Brown and A. B. Klots generously read and commented 

VOLUME 28, NUMBER 2 125 

on a previous draft of this paper. The ready willingness of W. A. Weber 
of the Colorado University Herbarium to make plant determinations 
is greatly appreciated. Conversations with J. M. Burns, C. L. Reming- 
ton, and W. B. Watt provided valuable insights. R. Chehey, J. F. Emmel, 
M. S. Fisher, K. Johnson, J. Scott, and A. O. Shields freely gave helpful 
field observations on Colias. Special thanks go to my wife Lydia Thomp- 
son, whose assistance in the field was invaluable. 


Ax, S. A. 1959. A study of hybrids in Colias (Lepidoptera, Pieridae). Evolution 13: 

Barnesy, R. 1964. Atlas of North American Astragalus. Mem. N.Y. Botan. Garden: 
13. 2 vol. im + 1188 p. 

Epwarps, W. H. 1873. Butterflies of North America. Vol. 1. Supplementary 
Notes. Amer. Entomol. Soc., Philadelphia. 215 p., 50 pls. 

1897. Butterflies of North America. Vol. III. Supplementary Notes. 
Houghton Mifflin, Boston. viii + 432 p., 51 pls. 

Enreuicu, P. & A. Exruicn. 1961. How to Know the Butterflies. Brown, Dubuque, 
Iowa. 253 p. 
Ferris, C. D. 1972. Notes on certain species of Colias (Lepidoptera: Pieridae) 
found in Wyoming and associated regions. Bull. Allyn Mus. Entomol. 5. 
1973. A revision of the Colias alexandra complex (Pieridae) aided by 
ultraviolet reflectance photography with designation of a new subspecies. 
J. Lepid. Soe. 27: 57-73. 

Hovanitz, W. 1950a. The biology of Colias butterflies. I. The distribution of 
the North American species. Wasmann J. Biol. 8: 49-74. 

. 1950b. The biology of Colias butterflies. II. Parallel geographical varia- 

tion of dimorphic color phases in North American species. Wasmann J. Biol. 

8: 197-219. 
Kuots, A. B. 1951. A Field Guide to the Butterflies. Houghton Mifflin, Boston. 
xvi + 349 p. 

Masters, J. H. 1970. Concerning Colias eurytheme alberta Bowman (Pieridae). 
J. Res. Lepid. 9: 97-99. 

McDunnoucu, J. 1922. Notes on the Lepidoptera of Alberta. Can. Entomol. 54: 

Suretps, O. A., J. F. Emmet & D. Breeptove. 1969. Butterfly larval foodplants 
records and a procedure for reporting foodplants. J. Res. Lepid. 8: 21-36. 




Biosystematics Research Institute, Canada Department of Agriculture, 
Ottawa, Ontario 

Coptodisca is a genus of very small moths. The forewing has silver 
and black markings on a white and yellow ground. The larva forms 
a mine in the leaf of its host by eating out the tissue between the 
upper and lower epidermis. When mature, the larva cuts a disc of 
tissue out of the leaf and uses it to form a cocoon. All known species 
of Coptodisca feed on woody plants, and most are restricted to a single 
plant genus. Nineteen species of Coptodisca have been described. 

Coptodiseca matheri Lafontaine, new species 
Pigs, eas 

Antenna fuscous, vertex of head golden, face and labial palps white. Thorax 
and abdomen silver-grey above, white below. Forewing with silvery white basal half, 
and. light yellow apical half. A (costal) spot two thirds of distance along costa 
and a spot opposite it on inner margin, each extending one third of the way across 
wing. Costal spot white, margined with dark grey. Spot on inner margin light 
grey, margined with dark grey. A grey patch at tornus which extends from distal 
edge of spot on inner margin to dorsum and termen. Apical patch wedge-shaped 
with a circular black base followed by a fan-shaped row of scales with light brown 
bases and black tips. Apical patch preceded and margined on both sides by white 
scales, separated from dark patch at tornus by yellow ground basally and some 
white scales distally. Yellow ground blending into dark patch at tornus between 
apex of spot on inner margin and apical patch. A small black wedge distal to 
costal spot, parallel to outer margin of spot. Cilia whitish grey, with black pencil 
of scales extending outward from apical patch. Hindwing and cilia uniformly 
whitish grey. Expanse 4.1 + .3* mm. (6 specimens). 

Male genitalia (Fig. 1): Genitalia excluding valves, 2% to 3 times as long 
as wide. Comb on valve with 6 teeth. Saccular setae on inner surface of valve 
in two small patches, one near dorsal edge and one near apex. 

Female genitalia (Fig. 3): Similar to those of other species of Coptodisca 
except for tip of ovipositor. Central point of five pointed ovipositor is smallest. 
In other species of Coptodisca teeding on plants in Ericaceae central point enlarged 
and bulb-shaped, much larger than the other four points. 

Type material: Holotype male, Jackson, Mississippi, emerged 7 September 1965 
(Bryant Mather). Reared from Vaccinium arboreum Marsh. Type No. 13032 in 
Canadian National Collection. Allotype female, Jackson, Mississippi, emerged 25 
January 1965 (Mather). Paratypes, one male, Jackson, Mississippi, emerged 25 
January 1965 (Mather); two males, Clinton, Mississippi, emerged 22 December 
1969 (Mather). Allotype and paratypes reared from same host as holotype. All 
specimens reared in laboratory at Ottawa. 

* standard deviation 


Figs. 1, la, 2. Male genitalia of Coptodisca spp.: 1, C. matheri n.sp.; la, right 
valve of C. matheri; 2, right valve of C. negligens Braun. 


Fig. 3. Female genitalia of Coptodisca matheri n.sp. 

Distribution: Known only from central Mississippi but undoubtedly more widely 

Foodplant. Farkleberry (Vaccinium arboreum Marsh. ) 

Mine. The mine usually follows the leaf margin. It begins at the base of 
the leaf near the top of the petiole as a serpentine mine, and widens into an 
elongate blotch slightly wider than the oval disc cocoon cut from the mine by the 
larva. The initial portion of the mine is completely filled with frass. Mr. Mather’s 
observations of the mining habits of this species on Farkleberry indicate that 
there are at least two generations each year. 

VoLUME 28, NUMBER 2 129 

Fig. 4. Coptodisca matheri n.sp., upperside of holotype. 

This is the most lightly coloured species of those which feed on 
plants of the Heath family. Coptodisca matheri can be separated from 
C. arbutiella Busck (1904: 769) and C. kalmiella Dietz (1921: 44) by 
the lack of the dark lead ground colour on the basal half of the forewing 
present in arbutiella and kalmiella. The comb on the valve of the male 
genitalia of matheri has 6 teeth, not 7 to 9 as in arbutiella and kalmiella. 

Coptodisca matheri is most easily separated from C. magnella Braun 
(1920: 79) by its lack of a sharp contrast on the forewing between 
the yellow ground colour and the dark patch at tornus. In matheri the 
pale yellow ground colour gradually blends into the dark colour of 
the patch at tornus. 

Coptodisca matheri differs from Dr. Braun’s (1916: 138) description 
of C. negligens, and from specimens which match her description reared 
from Vaccinium angustifolium Ait., by the lighter yellow colour on the 
forewing, and by the golden rather than grey colour of the scales on 
the vertex of the head. The valve of the male genitalia of matheri (Fig. 
la) is much less extensively setose than that of negligens (Fig. 2). The 
comb on the valve of matheri has 6 teeth not 5 as in negligens. 

The following is a key to the known species of Coptodisca which feed 
on species of plants in the Heath family (Ericaceae). 


1—Yellow ground colour continuous between costal spot and spot 

Oh inher Margin... 2 
—Ground colour interrupted between these spots by a mottling 
of dark scales... ee ee eee 4 
2—Head silvery-lead, concolorous with thorax and base of forewing; 
Arbutusteeder = 2.) 8 ieee C. arbutiella 
—Head golden 0 ee eee 3 
3—Sharp contrasting line where yellow ground meets dark posterior 
patch; (Gaylussacia-teeder, ===) = ar C. magnella 
—Yellow ground colour blending gradually into dark patch at 
tornus: Vaccinium-teeder 22 eee C. matheri 

4—Comb on valve of male genitalia with 7-9 teeth; Kalmia-feeder _ 
EpipE Wnt eer nmemea ome ULAR ONCE E Vin VEN Ne C. kalmiella 

—Comb on valve of male genitalia with 5 teeth; Vaccinium-feeder 
BAlnas ss) eine atenen i NE C. negligens 


I wish to thank Dr. T. N. Freeman formerly of the Entomology 
Research Institute, Canada Department of Agriculture, for reviewing 
this manuscript and making helpful suggestions. 


Braun, A. 1916. New species of Microlepidoptera. Can. Entomol. 48: 138-141. 
. 1920. New species of Lyonetiidae (Microlepidoptera). Entomol. News. 
31: 76-80. 
Buscx, A. 1904. Tineid moths from British Columbia, with descriptions of new 
species. Proc. U.S. Nat. Mus. 27: ‘745-778. 
Dietz, W. G. 1921. A new species of Coptodisca (Lepid.) Can. Entomol. 53: 44. 




Adult size and weight have been related to female fecundity for a 
number of Lepidoptera. Weight of emerging moths is positively corre- 
lated with numbers of eggs deposited for Ephestia elutella Hiibner 
(Waloff, Norris & Broadhead, 1948). Because weight may vary greatly 
with age and with environmental conditions such as temperature and 
humidity, more intrinsic and stable indicators of moth size are desirable. 
Wing length, a more constant indicator of moth size than body weight, 
has been used as an indicator of fecundity for Crambus harpipterus 
Dyar and Agriphila plumbifimbriella Dyar (Crawford, 1971), and for 
Oncopera intricata Walker (Martyn, 1965). 

Pupal size was used as an indicator of fecundity by Williams (1963) 
for Proceras sacchariphagus Bojer and by Miller (1957) for Choristo- 
neura fumiferana (Clemens). Johnson (1968) used the same approach 
as Miller to establish the relationship between pupal size and fecundity 
of C. pinus Freeman. In these instances, adult moths were reared from 
measured pupae and allowed to deposit their eggs. The amount of 
oviposition in turn was related to pupal size. This approach has the 
advantage that once the relationship between oviposition and pupal 
size has been determined, then fecundity can be estimated from empty 
pupal cases after moth emergence. Disadvantages include problems 
associated with rearing pupae to adulthood, and ovipositional per- 
formance of emerging females. 

A count of the developing odcytes in pupae approaching eclosion offers 
a possible index of potential fecundity. The present study was designed 
to determine if potential fecundity could be estimated from overwin- 
tering pupae of Rhyacionia neomexicana (Dyar). This paper relates 
oocyte complements of R. neomexicana pupae to two measures of pupal 


R. neomexicana overwinters as pupae enclosed within cocoons. Co- 
coons are attached to the root collars of host trees, Pinus ponderosa 
Laws., an average of 2.63 + 0.90 cm (n = 67) beneath the soil surface. 

1 Research Entomologist, U.S. Department of Agriculture, Forest Service, Rocky Mountain 
Forest and Range Experiment Station, with central headquarters maintained at Fort Collins in 
cooperation with Colorado State University; author is located at Albuquerque in cooperation 
with the University of New Mexico. 


Fig. 1. Female Rhyacionia neomexicana pupa showing two measures of pupal 
size: (a) linear distance from anterior edge of frontal horn to apex of right wing 
pad, and (b) width of 5th abdominal segment. 

A sample of overwintering pupae was dug 26 March 1970 about 10 days 
before initial male emergence and 20 days before initial female emer- 
gence. Cocoons were dug from the root collars of a natural stand of 
young pines on the Dudley Burn, Chevelon Ranger District, Coconino 
County, Sitgreaves National Forest, Arizona. They were placed in an 
ice cream carton with moist soil, transported to the laboratory (AI- 
buquerque) in an ice chest, and stored in a refrigerator (ca. 5°C) 
until dissected. 

In the laboratory, the cocoons were opened and pupae sexed by location 

VOLUME 28, NUMBER 2 133 




Total odcytes 

y =205.9x-156.4 



18 19 20 Z| 2.2 2.3 2.4 (215) 26 
Fifth abdominal segment width (mm) 

Fig. 2. Total odcytes of Rhyacionia neomexicana pupae as a function of abdominal 
segment width. 

and configuration of the genital pore. Live female pupae were mea- 
sured to the nearest 0.1 mm using a dissecting microscope equipped 
with an ocular micrometer. Two measurements were made on each 
pupa: (1) the linear distance from the anterior edge of the frontal 
horn to the apex of the right wing pad (Fig. la), and (2) the maximum 
width of the 5th abdominal segment (Fig. lb). The 5th segment is 
completely free of the wing pads, and remains intact after adult emer- 

Measured pupae were heat killed with a flamed dissecting needle 
and partially embedded in paraffin to facilitate dissection. Specimens 
were flooded with physiological saline, and the 8 chains of developing 
ova (ovarioles) removed. To minimize possible differences in odcyte 
complements due to age or time of development, all pupae (n = 20) 
were killed and dissected within a 3-day period (30 March-1 April). 
Ovarioles from the 1st 10 pupae dissected were stained with Grenacher’s 
Borax Carmine to differentiate ripe from unripe odcytes (Crawford, 
1971; Williams, 1963). For staining, ovarioles were submerged in the 
staining solution for 5 minutes, then de-stained by washing in 70% 
ethanol for 20-30 seconds. Ovarioles from the Ist 10 pupae were stained 



:3 250 
S A 
ie y=102.6x-237.5 
200 r=0.80 

46 47 48 49 50 5| 52 53 54 55 56 57 Se S9omcoma 
Frontal horn-wing tip length (mm) 

Fig. 3. Total odcytes of Rhyacionia neomexicana pupae as a function of frontal 
horn-wing tip length. 

and their odcytes counted on the day of dissection, while ovarioles from 
the 2nd group of 10 pupae were stored in 70% ethanol and counted 
(unstained) at a later date. Only discrete, differentiated odcytes were 
counted. Undifferentiated, developing odgonia in the germarium were 
not included. 

Regression analyses were run to determine the relationship between 
odcyte complement and 5th abdominal segment width and _ frontal 
horn-wing tip distance. The resulting equations were tested for the varia- 
tion in Y explained by the fitted line at Fo; with 4s df. 


All ovarioles (n = 80) in the Ist series of 10 pupae contained de- 
veloping odcytes that retained the stain, indicating the chorions im- 
pervious to the stain had not yet developed. Mean width of the 5th 
abdominal segment in dissected pupae was 2.18 + 0.20 mm, and mean 
frontal horn-wing tip distance was 5.17 + 0.40 mm. Total odcytes 
(5,869) had a calculated mean of 293.45 + 50.99 per pupa (range 

VOLUME 28, NUMBER 2 135 

Both regression equations (Figs. 2 and 3) were highly significant, 
P < 0.005. Thus, a reasonable estimate of o6cyte complement for over- 
wintering R. neomexicana pupae can be obtained from measures of 
pupal size. The equations indicate an increase of about 20-21 eggs per 
mm of abdominal width or frontal horn-wing tip length. 

The abdominal width measure may be more useful than the frontal 
horn-wing tip measure for estimates of potential fecundity. The 5th 
abdominal segment remains intact after adult emergence while the 
frontal horn-wing tip unit is ruptured and displaced during eclosion. 


Large overwintering R. neomexicana pupae as a rule have more 
odcytes in their ovarioles than do small pupae. Regression equations 
demonstrate the linear relationships between pupal size and numbers of 
oocytes found in the ovarioles. However, oocytes in the pupal stage 
must be considered only as. potential fecundity because additional 
odcytes may be differentiated after adult emergence, and some odcytes 
may be reabsorbed. Waloff et al. (1948) found that most of the eggs 
produced by Ephestis elutella Hiibner were present at the time of adult 
emergence, but that virgin females reabsorbed 40% and fertilized females 
reabsorbed 17% of their egg rudiments. An 11% reabsorption of unripe 
eggs in the ovaries has been reported for Proceras sacchariphagus Bojer 
(Williams, 1963). 

Other factors which may influence egg production are: availability 
of water and nutrients to emerging adults (Waloff et al., 1948; Williams, 
1963); density and nutrition of larval populations (Martyn, 1965; Miller, 
1957); mating condition of females, i.e., virgin vs. fertilized (Williams, 
1963); environmental effects on oviposition, survival, and longevity of 
females (Waloff et al., 1948); environmental effects on larval stages 
(Cook, 1961; Tantawy & Vetukhiv, 1960); and changes in the genetical 
constitution (Willington, 1964). These factors should be considered 
and explored before assigning a mean fecundity to a population. 


Coox, L. M. 1961. Influence of larval environment on adult size and fecundity 
in the moth Panaxia dominula L. Nature, Lond. 192 (4799): 282. 

Crawrorp, C. S. 1971. Comparative reproduction of Crambus harpipterus and 
Agriphila plumbifimbriella in Northern New Mexico. Ann. Entomol. Soc. Amer. 
64: 52-69. 

Jounson, S. A. 1968. Biotic environmental effects on the fecundity of jack pine 
budworm. MF Thesis, Univ. Mich. School of Natural Resources, 62 p. 

Martyn, E. J. 1965. Studies on the ecology of Oncopera intricata Walker 
(Lepidoptera: Hepialidae). I. Fecundity of the female moths. Aust. J. 
Zool. 13: 801-805. 


Miter, C. A. 1957. A technique for estimating the fecundity of natural popu- 
lations of the spruce budworm. Can. J. Zool. 35: 1-13. 

Tantawy, A. O. & M. O. Veruxuiv. 1960. Effects of size on fecundity, longevity 
and viability in populations of Drosophila pseudoobscura. Amer. Nat. 94: 

Watorr, N., M. J. Norris & E. C. BroapHEeap. 1948. Fecundity and longevity 
of Ephestia elutella Hiibner (Lep. Phycitidae). Trans. Roy. Entomol. Soc. 
Lond. 99: 245-267. 

WELLINGTON, W. G. 1964. Qualitative changes in populations in unstable environ- 
ments. Can. Entomol. 96: 436-451. 

WituiaMs, J. R. 1963. The reproduction and fecundity of the sugar cane stalk 
borer, Proceras sacchariphagus Bojer (Lep. Crambidae), p. 611-625. In J. R. 
Williams (ed.), Proceedings of llth Congress of the I.S.S.C.T., Mauritius, 
1962. Elsevier, Amsterdam. 


P.O. Box 475, Geraldton, Western Australia 6530 

The information in this paper has been extracted from notes recorded 
by the author between 1962 and 1964. As basic information on the 
larvae of H. n. nuttalli and H. h. hera apparently still remains to be 
published (Ferguson, 1971, p. 137-147), it seems worthwhile to publish 
these notes without further delay. The larval descriptions that follow 
remain valid, despite the passage of time, although it is quite con- 
ceivable that one or more of the localities mentioned has since been 
altered (perhaps even obliterated) by those activities of Homo sapiens 
popularly termed “development” and “progress.” 

Hemileuca (Pseudohazis) nuttalli nuttalli (Strecker ) 

In late April 1962, more than 100 completely-black and unmarked 
Hemileuca larvae (of the subgenus Pseudohazis), in various instars from 
quite small (second or third instar) to nearly fullgrown, were given 
to me by Ken Goeden, who collected them on 25 April 1962, near the 
highway in low hills between 11 to 13 mi. west of Vale, Malheur Co., 
Oregon (elevation about 2800 ft.). He found them resting and feeding 
on bitterbrush, Purshia tridentata (Pursh) DC. (Rosaceae), which was 

VOLUME 28, NUMBER 2 137 

growing there in an association with the abundant and widespread 
Great Basin sagebrush, Artemisia tridentata Nutt. (Asteraceae ). 

The living final instar larvae of this eastern Oregon population of 
nuttalli were briefly described (and readily recognized) as follows: 
Skin uniformly dull black, with NO maculation; a slight shading toward 
brown on the venter, especially in thoracic region. Body covered with 
a floccose pubescence of fine, soft, grayish-white hairs. No variation 
in body (or spine) coloration evident. Spines and spine clusters all jet 
black. The unbranched, short, sharp, clustered dorsal spines have a mild 
but definite urticating ability, if pushed firmly against tender skin. Head 
and thoracic legs blackish to brownish-black; faintly shiny. Head pubes- 
cent. Several of these larvae (code-numbered St.7), including the cor- 
responding notes, are preserved in my former North American larval 
collection (most of which now belongs to the Natural History Museum 
of Los Angeles County, California). 

As an interesting aside, concerning foodplant tolerances, I should men- 
tion that a number of the larvae collected by Goeden (1962) were also 
sent to Christopher Henne, at Pearblossom, Los Angeles Co., California. 
It was necessary for him to locate a substitute plant species, and as 
a natural first guess he offered them (the viscid) antelope brush, 
Purshia glandulosa Curran, which grows south and southeast of Pear- 
blossom (near Valyermo), but this plant was absolutely refused by the 
larvae! Next, Cercocarpus betuloides Nutt. ex T. & G. (also Rosaceae ) 
was offered; surprisingly, they readily accepted this substitute of another 
genus in preference to the other Purshia. Several of Henne’s larvae 
ultimately pupated and later produced perfect adults. I attempted 
to feed some of my captive larvae on Artemisia, but this was completely 
refused. The rest of my series died in the larval stage, due to lack of 
proper treatment in captivity. (See notes at end.) 

The adult moths (reared by Henne) emerged in the summers of 
1962 and 1964 (3 ¢ 4 between 12-28 Aug. 1962; one ? on 13 Aug. 1964). 
Showing only minor variation in colors and markings, they were briefly 
described as follows: Forewing upperside groundcolor dull but chalky 
whitish, sharply-marked with black; sometimes with a suffusion of 
yellow-orange at outer margin, just inside the narrow black border. 
Hindwing upperside rich yellow-orange, sharply marked with black. 
Undersides of both wings uniformly dull yellow-orange, sharply-marked 
with black. Thorax and abdomen yellow-orange, in some cases slightly 
marked with black. 

J. S. Buckett mentioned that he observed a diurnal flight of moths, 
fitting the above description, at around 1400 hrs. on 4 September 1963, 
in an area about 2 mi. W of Irrigon, Morrow Co., Oregon (elevation 


about 400 ft.). Many of the freshly-emerged adults were resting on 
Purshia bushes, and “thousands” were seen on the wing. At the time 
of this observation the temperature was about 90° F.; there had been 
a substantial rain in the locality some days before the emergence. It 
was noted that the moths were restricted to areas where Purshia was 
growing. Two of the adults (¢ and 2?) collected by Buckett are in 
my present collection; presumably others are in the Buckett collection. 

Hemileuca (Pseudohazis) eglanterina eglanterina (Boisduval) 

On 14 June 1964, David L. Mays gave me 40 gregarious first instar 
larvae of an unidentified Hemileuca sp. (subgenus Pseudohazis), which 
he had collected a few days earlier on Purshia tridentata (det. Mays), 
about 3 mi. N of Markleeville, Alpine Co., California, southeast of Lake 
Tahoe (at about 7500 ft. elevation). I transported these larvae south, 
to White Cliff Ranch, near Valyermo, Los Angeles Co., California, hoping 
to continue the rearing; there they readily accepted mature and semi- 
mature leaves of the local Cercocarpus betuloides as a substitute food- 
plant, and grew rapidly from second and third instars to maturity on 
that plant; this was followed by healthy pupae. (Incidentally, they also 
practically refused to accept the local Purshia glandulosa, which was of- 
fered as the first potential substitute, although a little feeding did take 
place on it.) 

The final instar larvae (20 July 1964), were briefly described as fol- 
lows: Dorsum contrasting abruptly with venter. Skin dull black down 
to the prominent, undulate cream-white subspiracular line; below this 
line, including prolegs completely, skin grayish-flesh-pink to pinkish- 
brown. Subspiracular line white, with two narrower whitish supra- 
spiracular lines; the uppermost lines much-suffused by large blotches 
of pale pinkish-purple centered between them, and blocking them out 
at intervals. Spines in the two dorsal rows of short, sharp (stinging) 
spine-clusters black, toward center of each cluster, but outermost spines 
of these clusters pale straw-yellow, minutely black-tipped. (In earlier 
instars, these identical spine-clusters contained predominantly light 
golden-brown spines.) The longer subdorsal and lateral spines primarily 
black. Body covered with fine, soft, grayish-white hairs. Head shiny 
blackish-brown with faint reddish-purple tinge; pubescent. Thoracic 
legs black and glossy. This description was drawn from notes on several 
of the living Markleeville larvae; some of these were preserved under 
my code-number St.16 (now in the Natural History Museum of Los 
Angeles County). It would not be surprising if larvae of this species, 
from various widely-separated populations, were found to show con- 
siderable variation in color and/or maculation. 

VOLUME 28, NUMBER 2 139 

These larvae were gregarious when small, clustering together both 
while resting and while feeding. They always followed each other in 
a perfect single file procession when moving to new locations. This 
gregarious behavior was gradually lost as they grew older, becoming 
essentially non-existent in last instar. 

All pupae obtained were given to Christopher Henne; adults (1 ¢; 
1 2) emerged in July 1965. Pupation took place in the typical 
“Hemileuca-type” of surface-debris cocoon or cell, under some sheltering 
object (such as a rock or board), but always on or only just below the 
soil surface; soil grains, small pebbles, and any other nearby particles 
of litter, were densely-incorporated into the relatively soft and flexible, 
silk-tied cell walls. 

Hemileuca (Pseudohazis) hera hera (Harris) 

For comparison with the larvae just described, a brief description of 
the final instar larva of Hemileuca hera (based on a few living individuals 
from one population) seems worthy of inclusion here. I collected these 
larvae on 26 July 1964, when on a brief trip with Edmund C. Jaeger, in 
the Inyo Mountains, Inyo Co., California, at approximately 10,000 ft. 
elevation, about 3 mi. SW of Waucoba Peak. They were feeding on 
Artemisia tridentata (Great Basin sagebrush), which was growing 
patchily in the more open areas of a bristlecone pine forest (Pinus 
aristata). On this date both penultimate and last instar larvae were 
present on their foodplant; unhatched egg-masses were also noticed, 
and a few females were observed in the act of ovipositing. Many adult 
males were on the wing; a few pairs were seen in copulation, resting 
on the sagebrush. 

When transported to White Cliff Ranch, near Valyermo in the San 
Gabriel Mts., (elevation close to 5000 ft.), these larvae hardly nibbled 
at the local sagebrush (probably a distinct subspecies or variety of A. 
tridentata), and soon began to decline as starvation ensued. None of 
them filled out or reached a prepupation condition. They were also 
offered Cercocarpus betuloides as a last resort, but this plant was totally 
refused. Daily sunlight and fresh air were provided, so it was not for 
lack of these that the larvae died. It is possible that the rapid drop 
in elevation had as much of a bad effect on them as did my attempt 
to force them onto a distinctly different form of the foodplant species; 
both of these factors were probably responsible for their decline. 

The final instar larvae (McFarland code-number St.17 in the Los 
Angeles County Museum), were briefly described as follows: Skin of 
dorsum and sides dull black; venter, including bases of prolegs, pale 
grayish-brown. Dorsum and sides marked with several full-length lines 


of cream-white: a closely-parallel pair of narrow, broken middorsal 
lines; a broader and nearly solid subdorsal line; a slightly narrower, 
undulate supraspiracular line; a similar undualte subspiracular line. 
Body covered with a fine, soft, grayish-white pubescence. Spines in the 
two dorsal rows of short, sharp (stinging) spine-clusters mostly pale 
straw-yellow basally and widely-tipped with black. Longer lateral spines 
primarily glossy black with some straw-colored basal branches. Thoracic 
legs and lateral shields of prolegs glossy black. Head deep glossy black; 

Eggs collected on Artemisia tridentata at the Inyo Co. locality, were 
briefly described as follows: deposited on the foodplant twigs, in com- 
pact, and securely-glued encircling-bands, with no covering of scales, 
“fluff,” or dried froth, etc. Among those egg “masses” observed, numbers 
ranged between 20 and 80 eggs per mass. The chorion was very tough, 
smooth, and glossy. Color at this stage (not long after oviposition) was 
a uniform pale whitish-gray-green, without bands, spots, or other macu- 
lation. The egg color was rather close to that of the (Inyo Mts.) sage- 
brush leaves, but had less green in it. The eggs probably overwinter, 
hatching perhaps in late May or sometimes in June at this elevation. 

H. hera, and its abundant (often dominant) widespread foodplant, 
Artemisia tridentata Nutt., are also present in eastern Oregon, but the 
Purshia-feeding H. nuttalli nuttalli appears to be of more localized 
distribution there, probably only occurring in certain areas where its 
(less-abundant) foodplant grows. 

Some reared adults from the above-described Hemileuca larvae, with 
the exception of hera, are in the Henne collection. 

Notes on Rearing Hemileuca Larvae Successfully in Captivity 

Most Hemileuca larvae can prove to be delicate in captivity, and will 
usually decline (slowly) and die IF deprived of fresh air and sunlight. 
Daily SUNLIGHT appears to be particularly important to stimulate 
vigorous feeding and normal, healthy growth in these larvae. (Electric 
lighting can be used but is only a poor substitute; never use “Cool White” 
fluorescent.) If housed in thoroughly-ventilated cages, with sprigs of 
foodplant kept fresh in water, and if given about one hour of sunlight 
daily (or at least as often as it is available), they will thrive and are 
definitely NOT difficult to rear. A light sprinkling of water over the 
foodplant, at least every second or third day (in the early morning), 
is highly desirable. With reference to the sun requirement, it is impera- 
tive that some shade also be available at all times during the sunning- 
period, so that the larvae can move quickly and easily from a sunny 
location into the shade as individually required. To provide such condi- 

VoLUME 28, NUMBER 2 14] 

tions, a cage that is all plywood on top and on two opposite or adjacent 
sides, with screen only on the other (two) sides, is ideal. This makes 
it possible to safely leave the cage in a completely sunny location all 
morning, without any need of further attention, while insuring that there 
will constantly be areas of both sun and shade within; ample ventilation 
is also provided. If it is semi-cloudy, it may sometimes be necessary 
to leave the cage in a potentially sunny location all morning in order to 
accumulate enough actual “sun-time” to benefit the larvae; yet, on a 
hot and clear morning, they might be urgently needing to seek shade 
within less than an hour after the sunning began. The constant presence 
of some zones of shade in the cage will also guarantee less drastic wilting 
of (at least a portion of ) the foodplant sprigs—another important factor 
in many cases. 

Incidentally, the above suggestions will also be found helpful in 
connection with a number of other “difficult” bombycoid larvae in 
captivity, such as the rare Californian saturniid, Saturnia albofasciata 
(Johnson), and certain Australian anthelids (some Pterolocera and 
Anthela spp.); also applicable to a few arctiids (some Apantesis spp.), 
many agaristids, and to a wide scattering of unrelated genera in various 
other macro families where strictly diurnal-feeding larvae are involved. 


I would like to thank D. S. Fletcher (British Museum Nat. Hist.) for 
recently reviewing this manuscript, and for thoughtfully providing needed 
photocopies of relevant pages from the reference cited below. I am 
deeply indebted to Ken Goeden (Oregon) for giving me the nuttalli 
larvae, to David L. Bauer (Calif.) for a most helpful discussion (letter: 
26 January 1966) further verifying the species described in this paper, 
and to Christopher Henne (Calif.) for completing the rearing of the 
Markleeville larvae when I was preparing to leave for Australia in 1964. 


Fercuson, D. C. in Dominick, R. B., et al., 1971. The Moths of America North 
of Mexico, Fascicle 20.2A, Bombycoidea (in part). Classey, London. 153 p., 
11 color plates, 19 figs. 



P.O. Box 5111, APO New York, 09286 

Ferguson's treatment of the Saturniidae (1971) provided much valu- 
able information for those particualrly interested in Hemileuca. He 
presented certain questions for further study, as he had not enough 
material on hand to make definite statements. I want to record here 
some of my experiences in rearing members of this group and provide 
information that might assist in clarifying some of the shadowy areas. 
I also would like to add some of my rearing methods and misfortunes 
in hopes that they will help others to avoid my errors and thus be suc- 
cessful in their initial attempts at rearing the members of this beautiful 


Hemileuca maia (Drury) 

Ferguson mentions the great confusion that exists between the oak 
eating H. maia and the willow eating H. nevadensis (Stretch), including 
the lack of information available to verify the acceptability of foodplants 
other than Quercus for maia. In May and June 1972 I reared maia from 
ova received from Irwin Leeuw of Cary, Illinois. These ova were col- 
lected on scrub oak at Colonie, Albany Co., New York, on 15 April 
1972. This is well within the range of true maia and well away from 
the influence of nevadensis. I successfully reared these larvae to ma- 
turity on Salix (willow) from the Mojave riverbed near Victorville, 
San Bernardino Co., Calif. I tentatively identified the willow as sandbar 
willow, (Salix hindsiana Benth). The larva readily accepted this as an 
alternate foodplant after feeding on a California scrub oak (Quercus 
chrysolepis Liebm.) for two instars. I had an 80% successful pupation 
rate and emergence began in September 1972. So apparently maia does 
accept Salix, at least in captivity. I leave it to the Midwest collectors 
to solve the maia-nevadensis confusion in that area. 

Hemileuca electra (W. G. Wright) 

Ferguson mentions that he saw too few specimens of H. electra clio 
(Barnes and McDunnough) to give a definite statement on the validity 
of its status as a subspecies. Southern California collectors who have 
had experience with this species feel that clio extends its range into 
California on the Mojave Desert plateau to the desert foothills of the 

VOLUME 28, NUMBER 2 143 

San Bernardino, San Gabriel, and Sierra Nevada mountains. Larvae 
and ova masses are found on Eriogonum fasciculatum var. poliofolium 
(Benth) within this range and the adults match closely to clio. H. 
electra electra on the other hand occurs on the coastal slopes of these 
ranges, to the ocean, feeding on nominate E. fasciculatum (Benth). In 
the 1972 season, I reared the larvae of these two subspecies side by 
side to find out if there were larval differences that might strengthen 
the validity of these two forms as subspecies. 

I took 20 first instar larvae of e. clio on 27 February 1972 at Rock 
Corral, 20 miles east of Lucerne Valley, San Bernardino Co., California. 
I also took 30, third to fifth instar, larvae of e. electra on 25 March 1972 
from one mile west of Lake Mathews Dam, Riverside Co., California. 
The following differences were observed in the physical appearances 
of the fifth instar larvae of each group. 

a. The spines on the lateral rows of e. electra were as described by 
Ferguson, “black with yellowish tips.” This trait was consistent on all 
the Lake Mathews larvae. All e. clio larvae from Rock Corral had the 
lateral rows of spines colored solid black, without any yellow tips. 

b. When compared, the e. clio larvae had much less white mottling 
or spotting on the body than did the e. electra, a characteristic that gave 
the e. clio larvae a much darker over-all appearance. 

c. The whitish line that flows lengthwise along the body of the larvae 
just above the spiracles is much more pronounced or “striking” in e. clio 
than in e. electra, and much straighter. 

These larvae all pupated in late April and early May 1972 and began 
emerging in July 1972. I have taken H. e. clio larvae or ova from the 
Rock Corral spot, from the foothills south of Apple Valley, San Bernar- 
dino Co., and from one mile north of Red Rock Canyon, off Hwy 14, 
Kern Co., all in California. 

After two years of unsuccessful attempts at rearing e. clio on its 
native foodplant, a very dry form of E. fasciculatum, in 1972 I transferred 
them to the nominate E. fasciculatum that e. electra feeds on. It is 
much longer-leafed and lusher, and I was successful in bringing the 
majority of the larvae through on this plant. 

Hemileuca burnsi (J. H. Watson) 

I have found larvae of this species commonly on Tetradymia axillaris 
(A. Nels) cotton thorn, and Prunus fasciculata (Gray) desert almond, 
in the foothills south of the Victorville-Apple Valley area of San 
Bernardino Co., California. These larvae are best collected in late 
January and early February when the foodplants are just beginning 
their growth and the black larval masses are easily spotted. Some 


collectors are successful in finding the oval rings in the winter, but 
I am not one of them. Where I have searched long and hard for ova, 
I have found many larval masses in the spring. I have had pupae from 
H. burnsi continue to emerge for two years after pupation. These larvae 
and those of the other Hemileuca that I have had experience with are 
very susceptible to parasites, and therefore are best taken in the earlier 

I add some general comments on my experiences with the rearing of 
Hemileuca. I have found that all the attitudes about them being hard 
to rear are true, and only after many unsuccessful attempts have I been 
able to bring a good series of adults out. I have found that the larvae 
require absolute cleanliness and constant, fresh food. They also cannot 
be crowded, and I limit them to 10 larvae per container in the fifth 
instar. I use clear plastic quart jars that can be purchased inexpensively 
in any store, and drill holes in the bottom for the stems of the food- 
plant to be put into water. The larvae are very susceptible to disease, 
and several can be lost in a short time. I have reared all my larvae with 
no sunlight, but with abundant artificial light. When the larvae begin 
to roam about the bottom of the rearing container and take on a 
discolored appearance, I transfer them to another container for pupa- 
tion. I use common “cat litter” as a pupating medium for all larvae, 
with tissues shredded on the top. This material seems to make very 
good pupal cells and is very mold resistant. The larvae usually burrow 
under the surface after a couple of days of roaming and pupate using 
the tissues as the top of the cell. The larvae frequently tend to pupate 
in groups, or near branches or twigs in the container. I wait two 
weeks after the last larva has burrowed before I gather the pupae; this 
allows enough time for all to pupate. Strangely, almost all the species 
in captivity begin emerging in July, although their natural flying period 
is September to November. This emergence continues off and on through 

In summary, my rearing experiences with several Hemileuca species 
have supplied the following data to help answer questions in shadowy 
areas: H. maia will readily accept Salix as an alternate foodplant, at 
least in captivity; H. e. electra and H. e. clio have definite, consistent 
larval differences that support the idea of subspeciation, and the range 
of H. e. clio extends into the northern desert areas of southern California; 
and H. burnsi will accept Prunus in the wild or in the laboratory. I hope 
these data will aid in clarification of the status of the species and sub- 
species of this beautiful group and encourage others to rear the larvae. 

VoLUME 28, NUMBER 2 145 

With Ferguson’s outstanding book for guidance, much more can be 
learned about Hemileuca through rearing and experimentation. 

I wish to extend my sincere thanks to Christopher Henne, of Pear- 
blossom, California, for his encouragement and eduction in life history 
work, and his kind review of this paper. 


Frercuson, D. C. in Dominick, R. B. et al., 1971. The Moths of America North of 
Mexico. Fasc. 20.2A, Bombycoidea, (in part), Classey, London, p. 101-153. 



Department of Zoology, University of Massachusetts, 
Amherst, Massachusetts 01002 

The relative dearth of information on the incidence of melanism in 
North American moths has been recently noted (Kettlewell, 1973). 
Since the reviews of Owen (1961, 1962) called attention to increasing 
melanism in various bark-like noctuids and geometers, little else on 
North American species has been published. Owen & Adams (1963) 
analysed the occurence of melanism in Catocala ilia (Noctuidae) in 
Michigan, and Klots (1964, 1966, 1968a, b) briefly noted increases in 
the frequencies of the melanic forms of Charadra deridens and Panthea 
furcilla (Noctuidae) in Connecticut. More recently, Sargent (1971) 
provided data on melanism in Phigalia titea (Geometridae) in central 
Massachusetts. The present data, acquired in the course of collecting 
moths for other studies in central Massachusetts from 1968-1973, are 
presented in hopes of stimulating others to acquire and publish similar 
data. Accumulated records, from different areas and at different times, 
may permit some meaningful geographic and _ historical comparisons, 
and so may contribute eventually to a thorough analysis of melanism 
in North America. Certainly every effort should be made to take 
advantage of our opportunity to study this phenomenon as it unfolds, for 
this opportunity may now be lost elsewhere in the world (Kettlewell, 


TasLeE 1. The numbers of typical and melanic individuals of Panthea furcilla 
taken in Leverett, Massachusetts (1970-1973). 


Forms 1970 1971 1972 1973 Totals 
Typical 29 AT 28 43 147 
Melanic 35 94 40 ri; 249, 
% Melanic 54.7 66.7 58.8 62.9 62.2 


The records included here involve only those species which have been 
substantially sampled, as my experience indicates that general impres- 
sions from limited samples are unreliable as indicators of melanic 
frequencies. The six species on which I report were collected in toto 
over their entire flight seasons during the years indicated, and virtually 
all of the specimens have been retained in my collection. 

Five of the six species considered were taken exclusively at light 
sources (incandescent, flourescent black light, and mercury vapor). 
Most of these specimens were obtained in a Robinson trap (mercury 
vapor) which operated from dusk to dawn, and, as expected, most of 
the specimens from light sources were males. One species, Catocala 
ultronia, was taken at both lights and bait, but there were no differences 
between the two samples, or between the sexes in the bait sample, in 
terms of melanic frequencies in this case. 

All specimens, unless otherwise indicated, were taken at my home in 
Leverett, Massachusetts. This collecting site is located in an extensive 
mixed deciduous woodland, most of which has grown up since a logging 
operation about 30-35 years ago. The dominant trees are oaks (Quercus 
velutina and Q. alba), with substantial representation of birches (Betula 
papyrifera and B. lenta), hickories (Carya glabra and C. ovata), pine 
(Pinus strobis), and hemlock (Tsuga canadensis). Some of the nearby 
area is more recently abandoned pasture, and is now in an intermediate 
stage of succession (sweet fern, Comptonia peregrina, juniper, Juniperus 
virginiana; gray birch, Betula populifolia; etc.). 

Leverett is located some 75 air-miles west of Boston, 25 air-miles 
north of Springfield, and 66 air-miles east of Albany, New York. The 
collecting area shows little visible evidence of air-borne pollution, as 
lichens abound on tree trunks which are not noticeably darkened by 
soot. I have previously referred to the area as “ostensibly rural” (Sar- 
gent, 1971), in an attempt to give recognition to both its visible ap- 
pearance and its location in the heavily industrialized northeastern 
United States. 

VoLUME 28, NUMBER 2 147 

TABLE 2. The numbers of non-melanic and melanic individuals of Catocala ultronia 
taken in Leverett, Massachusetts (1968-1973). 

Forms 1968 1969 1970 1971 1972 1973 Totals 
Non-melanic Al 22, 163 192 20 32 A470 
Melanic ri 7 Dp AT 5 10 98 
% Melanic 14.6 ANI 11.9 19.7 20.0 23.8 Wes 

Panthea furcilla (Packard). The melanic form of this species, 
atrescens McDunnough, is easily distinguished from its typical counter- 
part by the black ground of the wings, though melanics do vary con- 
siderably in the extent of their white lines (see figures in Ginevan, 1971). 
The genetic basis of melanism has been studied (Ginevan, 1971), but 
further work is required, particularly to determine whether heterozygote 
and homozygote melanic males can be distinguished reliably by visual 
inspection. The numbers of typical and melanic individuals taken in 
Leverett from 1970-1973 are presented in Table 1. 

Catocala ultronia Hubner. This highly polymorphic species has a 
strongly melanic form, nigrescens Cassino, with uniform, deep black 
forewings. This melanic was illustrated in Cassino’s paper (Lepidopterist 
1: 79, pl. vi), but is not shown in more popular works, such as Barnes 
& McDunnough (1918). The most common form of this species in 
Leverett is celia Hy. Edwards (Barnes & McDunnough, 1918: pl. VI, 
18), but all of the non-melanic forms are considered together in the 
tabulation of collecting results (Table 2). 

Catocala connubialis (Guenee). This generally rare moth has a 
“partly melanic’ form, pulverulenta Brower, with nearly uniform grayish 
forewings; and a strongly melanic form, broweri Muller, with uniform, 
deep green-black forewings (see figures in Muller, 1960). Most of the 

TABLE 3. The numbers of individuals of each form of Catocala connubialis taken 
in central Massachusetts (1970-1973). 

Forms Leverett West Hatfield Totals (%) 
sancta 1 — IL (C8335) 
cordelia 2 3 lier) 
pulverulenta 4 9 13 (43.3) 
broweri 3 8 ie ( SGA) 


TABLE 4. The numbers of typical and melanic individuals of Nacophora quernaria 
taken in Leverett, Massachusetts (1971-1973). 

Forms 1971 1972 1973 Totals 
Typical 3 10 iL 24 
Melanic 3 12 ive Be 
% Melanic 50.0 54.5 60.7 if a 

non-melanic specimens taken in this area are similar to, though darker 
than, cordelia Hy. Edwards (Barnes & McDunnough, 1918: pl. IX, 19), 
and occasional specimens are close to sancta Hulst (Barnes & McDun- 
nough, 1918: pl. IX, 21). Due to the rarity of this species, the numbers 
of specimens of the various forms are summed for the years 1970-1973, 
and I have included specimens taken in a Robinson trap at West Hat- 
field, Massachusetts (7.5 air-miles from Leverett) by Charles G. Kellogg 
(Malle 3): 


Nacophora quernaria (Abbot & Smith). The melanic form of this 
species, atresecens Hulst, is jet black, with only occasional traces of faint 
whitish along the ordinary lines. Specimens splotched with white on 
a blackish ground were considered typical, as were all brownish speci- 
mens. This species is generally uncommon in Leverett, but the frequency 
of melanic individuals has been consistent (Table 4). 

Biston cognataria (Guenée). Typical specimens of this species in 
Leverett are rather dark gray, being close to the insularia* category 
of Biston betularia in England (Kettlewell, 1973: pl. 9.1, no. 2, left). 
The melanic form, swettaria Barnes & McDunnough, is nonetheless 
easily distinguished, being uniformly black over the entire wing surfaces 
(Kettlewell, 1973: pl. 9.1, no. 3, right). The numbers of typical and 
melanic specimens taken in Leverett from 1971-1973 are presented in 

Table 5. 

TABLE 5. The numbers of typical and melanic individuals of Biston cognataria 
taken in Leverett, Massachusetts (1971-1973). 

Forms 1971 1972 1973 Totals 
Typical 23 22 84 129 
Melanic = 1 5 6 

% Melanic - 4.3 5.6 4.4 

VOLUME 28, NUMBER 2 149 

TasiE 6. The numbers of typical and melanic individuals of Phigalia titea taken 
in Leverett, Massachusetts (1968-1973). 

Forms 1968 #1969 1970 1971 1972 1973 Totals 
Typical 125 Es Silt 189 Jy 123 820 
Melanic 44 26 30 Al 32 34 207 
% Melanic 26.0 JG 18.6 ieee as), DLE 7 20.2 

Phigalia titea (Cramer). The records presented here (Table 6) will 
up-date those previously reported for Leverett (Sargent, 1971). The 
typical and melanic form, deplorens Franclemont, of this species are 
illustrated in Remington (1958). Melanism in this species is very 
clear-cut; well over 1000 specimens have been taken, and only one or 
two of these were difficult to assign to either the typical or melanic 


All of the species considered here presumably show industrial mela- 
nism, in the broad sense of that phrase. The melanics in these cases 
were extremely rare or absent in collections made prior to 1930 or 
1940, and now they comprise substantial proportions of the existing 
populations. However, the generally held explanation of industrial mela- 
nism, as developed by Kettlewell through studies on Biston betularia 
in England (Kettlewell, 1958), seems not completely applicable to the 
present results. This explanation stresses the cryptic advantage of me- 
lanics on darkened tree trunks, but the trees in the Leverett study area 
are not noticeably devoid of lichens or blackened by soot. Many of 
the melanics taken there are extremely dark, nearly jet black, and would 
seem to be cryptic on only the darkest trees in heavily polluted areas. 
Furthermore, the apparent tendency of some of these melanics to prefer 
light backgrounds, like their typical counterparts (Sargent, 1969), makes 
an explanation for their occurrence based on cryptic advantage even 
less likely. 

It is interesting to note that the frequency of melanics in Biston 
cognataria is quite low in Leverett, much lower, for example, than that 
occurring in New Haven, Connecticut (C. L. Remington, pers. comm. ), 
or in the areas in Michigan sampled by Owen (1961). Perhaps Biston 
spp. are industrial melanics in much the sense that Kettlewell has pro- 
posed (1958), but recent data cast some doubt on the completeness of 
a cryptic advantage explanation for even B. betularia (Bishop, 1972; 
Lees, Creed & Duckett, 1973). 


An activity of man which may have resulted in certain darkened 
backgrounds, and thus had an influence on at least one of the species 
considered here, is logging. This activity has been carried out periodi- 
cally over most of New England since Colonial days, and one of the 
most prized timber trees throughout this period has been white pine, 
Pinus strobis, the foodplant of Panthea furcilla. If this moth tends to 
rest on the trunks of this tree, then a tendency of loggers to take the 
larger trees (with lighter, furrowed bark), and leave the younger trees 
(with darker, smooth bark), may have provided an ecological oppor- 
tunity, in the form of more appropriate resting substrates, for melanic 

Certain other species, notably Phigalia titea and Nacophora quernaria, 
which have rather high melanic frequencies in Leverett, suggest that 
various factors associated with industrialization, other than observable 
environmental darkening, should be investigated with respect to the 
incidence of melanism. For example, air pollution affects the physical 
and chemical characteristics of vegetation, and perhaps the larvae of 
melanics are better able than the larvae of typicals to tolerate such 
changes. Certainly, melanics have exhibited superior viability in a 
number of physiological tests (Ford, 1937, 1940). Industrialization has 
undoubtedly also had deleterious effects on the predators of insects, and 
perhaps relaxed selection pressures have allowed melanics to survive 
where they previously could not have survived. 

All of the species considered here are bark-like cryptic species, and 
melanism in moths, particularly that associated with industrialization, 
has been largely restricted to such species. It is also generally true that 
melanism in those species studied is controlled by a single gene, with 
the allele for black being dominant to that for pale or typical coloration. 
These observations, together with our knowledge that the frequency of 
melanics may increase rapidly in a population, encourage some highly 
speculative ideas, which I will discuss very briefly. 

Perhaps many bark-like species have been exposed throughout their 
histories to recurring situations where melanism has been advantageous. 
If so, these species may have evolved mechanisms which enable them 
to change quickly from prevailingly pale to prevailingly dark popula- 
tions. Such changes might be effected through conditional genes for 
melanism, i.e. genes which are expressed only under conditions that 
are associated with environmental darkening. Among such conditions 
might be the chemical or physical effects of forest fires on the insects 
or their foodplants. The ability of a species to respond to these effects 
by producing adult melanics, which would then be cryptic on blackened 
backgrounds, could give it a clear selective advantage. Perhaps then, 


industrialization is creating conditions which are similar, or identical, 
to conditions created historically by forest fires. Melanic forms, in 
this event, would be somewhat analagous to the various seasonal forms 
which characterize many lepidopteran species. 

Much of this discussion is clearly fanciful speculation, but perhaps 
some excesses of this sort may be excused, if the result is to suggest that 
our understanding of industrial melanism is far from complete. 


Six species of bark-like moths with melanic forms were extensively 
sampled in central Massachusetts between 1968 and 1973. These species, 
and the percentages of melanic individuals in the sampled populations, 
are: Panthea furcilla (62.2%), Catocala ultronia (17.3%), Catocala con- 
nubialis (36.7%), Nacophora quernaria (57.1%), Biston cognataria 
(4.4%), and Phigalia titea (20.2%). These results are discussed with 
reference to various theoretical and speculative views on the phenomenon 
of industrial melanism. 


I thank Charles G. Kellogg for allowing me to use his unpublished 
data on Catocala connubialis. ; 


Barnes, W. & J. McDunnoucu. 1918. Illustrations of the North American species 
of the genus Catocala. Mem. Amer. Mus. Nat. Hist. 3, pt. 1. 

BisHop, J. A. 1972. An experimental study of the cline of industrial melanism in 
Biston betularia (1...) (Lepidoptera) between urban Liverpool and rural North 
Wales. J. Anim. Ecol. 41: 209-243. 

Forp, E. B. 1937. Problems of heredity in the Lepidoptera. Biol. Rev. 12: 461-503. 

. 1940. Genetic research in the Lepidoptera. Ann. Eugen., Lond. 10: 227- 

GinEvANn, M. E. 1971. Genetic control of melanism in Panthea furcilla (Packard) 
(Lepidoptera: Noctuidae). J. N. Y. Entomol. Soc. 79: 195-200. 

KETTLEWELL, H. B. D. 1958. Industrial melanism in the Lepidoptera and _ its 
contribution to our knowledge of evolution. Proc. 10th Int. Congr. Entomol. 
(1956) 2: 831-841. 

1973. The Evolution of Melanism, The Study of a Recurring Necessity, 
With Special Reference to Industrial Melanism in the Lepidoptera. Clarendon, 
Oxford. xxiv + 424 p. 

Krors, A. B. 1964. Notes on melanism in some Connecticut moths. J. N. Y. 
Entomol. Soc. 72: 142-144. 

1966. Melanism in Connecticut Panthea furcilla (Packard) (Lepidoptera: 

Noctuidae). J. N. Y. Entomol. Soc. 74: 95-100. 

. 1968a. Melanism in Connecticut Charadra deridens (Guenée) ( Lepidoptera: 

Noctuidae). J. N. Y. Entomol. Soc. 76: 58-59. 

. 1968b. Further notes on melanism in Connecticut Panthea furcilla 

(Packard) (Lepidoptera: Noctuidae). J. N. Y. Entomol. Soc. 76: 92-95. 


Lees, D. R., E. R. Creep & J. G. Ducxerr. 1973. Atmospheric pollution and 
industrial melanism. Heredity 30: 227-232. 

Mutter, J. 1960. A new melanic form of Catocala connubialis from New Jersey 
GNocemdaene J. epid Soe, 1A. aii lais: 

Owen, D. F. 1961. Industrial melanism in North American moths. Amer, Nat. 
95: 227-233. 

1962. The evolution of melanism in six species of North American 

geometrid moths. Ann. Entomol. Soc. Amer. 55: 695-703. 

& M. S. Apvams. 1963. The evolution of melanism in a population of 
Catocala ilia (Noctuidae). J. Lepid. Soc. 17: 159-162. 

Remincton, C. L. 1958. Genetics of populations of Lepidoptera. Proc. 10th Int. 
Congr. Entomol. (1956) 2: 787-805. 

SARGENT, T. D. 1969. Background selections of the pale and melanic forms of the 
cryptic moth, Phigalia titea (Cramer). Nature, Lond. 222: 585-586. 

1971. Melanism in Phigalia titea (Cramer) (Lepidoptera: Geometridae). 

J. N. Y. Entomol. Soc. 79; 122-129. 


PD. H. Haseck2 R. TI) Ansocasr® AN» iD) Girne 

Department of Entomology and Nematology, 
University of Florida, Gainesville, Florida 32601 

Schinia mitis (Grote) occurs from central Florida, north to Georgia, 
and west to eastern Texas (Hardwick, 1958). Most of the specimens 
Hardwick examined were collected in April, May, and June, but a 
few were collected in September and November. Kimball (1965) listed 
Florida records from March to June. Forbes (1954) gave the foodplant 
as Sitilias caroliniana Walt. [= Carolina false dandelion, Pyrrhopappus 
carolinianus (Walt.)DC]. Hardwick (1958) figured the lateral aspect 
of the egg and design of the chorion and gave a description and 
dimensions based on eggs dissected from preserved or dried females. 
Ganyard & Brady (1972) reported that males were attracted to virgin 
females of Indian meal moth, Plodia interpunctella (Hubner); almond 
moth, Cadra cautella (Walker); and fall armyworm, Spodoptera 
frugiperda (J. E. Smith), in field studies at Watkinsville, Georgia. No 
other published information was found on this species. 

1 Florida Agricultural Experiment Station Journal Series No. 5035. Received for publication. 

2 Research Associate, Florida State Collection of Arthropods. 

3 Present address: USDA, Stored-Product Insects R & D Laboratory, Box 5125, Savannah, 
Georgia 31403. 

VOLUME 28, NUMBER 2 153 

Fig. 1. Schinia mitis egg on inside of involucre of Pyrrhopappus carolinianus 
( Walt.) DC. 

Schinia mitis is a day-flying heliothidine moth. It is active only for 
a few hours in the morning when the flowers of the Carolina false 
dandelion are open. Moths fly rapidly from flower to flower but 
are easily netted while on a flower. On hot sunny days, all the 
flowers may be closed by 1000, whereas on cooler cloudy days, flowers 
may remain open until about noon. When the flowers are closed the 
moths usually rest facing downward on the stem or side of the receptacle. 
Occasionally, a moth was observed resting head downward in a flower. 
Mating apparently occurs on open flowers since mating pairs were fre- 
quently observed there from about 0700 to 0945, although mating 
probably also occurs earlier. Ovipositing females extend their abdomen 
downward between the florets and deposit eggs either on the developing 
ovaries or more often on the inside of the involucre (Fig. 1). 

During 1967, flowers of the Carolina false dandelion began to appear 
in late February in the Gainesville area; and a few blooms were observed 
as late as October. Most flowers appeared between mid-April and mid- 
June. Flowers collected on 18 and 24 March contained no larvae, but 
some eggs were found in the latter collection. Thereafter weekly col- 
lections consisting of 150 flower heads (50 open, 50 closed yellow, and 
50 closed white) were made through 11 August, and all samples con- 


TaBLE 1. Seasonal distribution and abundance of Schinia mitis eggs and larvae 
in weekly collections of flowers of Carolina false dandelion. Gainesville, Florida. 

50 Open 50 Yellow 50 White 
Flowers Closed Flowers Closed Flowers 

Date Eggs Larvae Eggs Larvae Eggs Larvae 
III-31 ils) 0 1155 2, 3) 14 
IV- 7 AB) 3 19 4 0 12 
-14 5 2 1 0 a 6 
-21 it 2, 2; 12 0 AS} 
-28 29 0 U5) 3 0) 19 
V- 5 30 1 19 28 0 38 
-12 6 4 4 17 0 31 
-19 19 3 ll: a 0 8 
-26 36 8) 2, 5 0 30 
VI- 2 7 ii 6 8 0 30 
- 9 2 0 4 A 0 20 
-16 19 1 15 2 0 12 
-23 6 0 13 9 0 21 
-30 16 0 1 3 0 i 
WAUls: 7 7 2 2 il 0 15 
-14 0 0 DZ 0 0 4 
-21 0) 0 0 1 0 2 
-28 0 0 0 0 0 4 
VIII- 4 23 0 17 i 0 Zit 
-ll 0 0 1 0 0 8 
Totals 246 I, evel 108 4 299 

tained eggs and/or larvae of S. mitis (Table 1). Flowers became scarce 
after that and partial samples collected on 18 and 30 August and on 
292, September were negative. This agrees with the results of Ganyard 
& Brady (1972) at Watkinsville, Georgia (ca. 300 mi NNW of the 
Gainesville area) where no S. mitis males were collected in traps placed 
in the field after about 10 September. 

Open flower heads contained mostly eggs, but almost no eggs were 
found in the closed white heads, although larvae were common there. 
In the yellow closed heads, both eggs and larvae were commonly found. 
This is not surprising in view of the development of both the insect 
and the flower. Sixty flower buds were tagged and checked daily to 
determine the flowering period. Each flower was open only 2-3 days 
followed by a closed period during which the seeds developed. During 
this closed period, the flower heads could be separated into a yellow 
phase (1.5-3.0 days) and a white phase (4.0-7.5 days) after which 
the head opened up and the seeds were blown away. Therefore, since 
oviposition cannot occur until the flowers open and the eggs do not 

VoLUME 28, NUMBER 2 ys 

Fig. 2. Pupa and larva of Schinia mitis: A, B, C. ventral, dorsal and _ lateral 
view of female pupa; D. lateral view of mature larva. 

hatch until the flower closes, there are only about 10 days in which 
the flower head is available for food. Since larval growth required a 
minimum of 13 days, the larvae must inhabit at least two flowers to 
complete their development. Large larvae were occasionally observed 
in open flowers or yellow closed flowers in the field. 

The sex ratio of 43 moths collected from flowers between 10 May 
and 30 June was nearly 1:1 (21 ¢, 22 2). Of 19 females examined, all 
had mated: 14 once, 4 twice and 1 three times. Four of the females 
contained over 40 fully developed eggs, with one having 75. 


On 2 occasions vespid wasps were observed burrowing into flower 
heads. On each occasion, when the wasp withdrew it was dragging a 
mature larva which it promptly stung. One of the wasps was captured 
and identified as Pterocheilus texanus Cresson. Predation by these and 
possibly other wasps apparently was not unusual since flowers were seen 
frequently that had been torn open in a similar way. The only other 
enemy observed was a crab spider, Mimusops sp., which had captured 
an adult moth on an open flower. 

Life History 

Eggs were collected from field flowers and placed singly in test tubes 
with whole or partial flowers and kept at 25° C and 14:10 light:dark 
photoperiod. All field collected eggs hatched within 3 days. Although 
24 larvae were reared to the pupal stage, only 18 emerged as adults, the 
others apparently entering diapause. The larval stage required 13-18 
days (avg. 15.3) and the pupal stage 9-17 days (avg. 12.9). Daily 
observations of 11 individuals revealed eight with 4 larval instars and 
three with 5 instars. The average duration of the stadia was 3.4, 2.4, 
2.4, and 7.0 for 4 instars and 4.3, 2.0, 2.0, 2.0 and 7.0 days for 5 instars. 
Development from hatching to adult emergence required 22-32 days 
(avg. 28.0). 

Egg (Fig. 1). Light green, closely matching color of inside of involucre. Color 
fairly constant at least until larva inside becomes visible. Dimensions of egg: 
length 0.71 + 0.03 mm, width 0.48 + 0.03 mm (23 eggs). 

Larva. Spicules, which become progressively more conspicious in later instars, 
present on bodies of all larvae. Spicules pale except in reddish transverse bands 
where they are black. 

First Instar. Head capsule width: 0.39 + 0.03 mm (n = 29). Head pale 
yellowish-brown. Ocellar area dark brown. Body yellowish-white. A _ faint 
yellowish-pink transverse band present on metathorax and abdominal segments 
1-8 in some specimens. Prothoracic shield, thoracic legs, anal shield, lateral 
sclerites on prolegs, and pinacula grayish-brown. 

Second Instar. Head capsule width: 0.65 + 0.04 mm (n = 28). Head 
light brown suffused with slightly darker markings. Ocellar area dark brown. 
Frons and adfrontals often lighter. Body pale orange except for whitish prothorax. 
Whitish spot dorsad and ventrad of seta D2 on abdominal segments. Prothoracic 
shield dark brown with 4 irregular incompletely separated black spots. Thoracic 
legs and pinacula dark brown. Anal shield and lateral sclerites on prolegs grayish- 

Third Instar. Head capsule width: 1.05 + 0.06 mm (n = 29). Head 
yellowish-brown, suffused with light brown markings. Ocellar area dark brown 
to black. Body reddish-brown, except lateral aspects of 10th abdominal segment 
whitish. Each abdominal segment with whitish spot dorsad and ventrad of 
seta D2. Metathorax with similar whitish spots plus a whitish subspiracular 
spot which is also present on abdominal segments 1 and 2. On mesothorax a 
whitish spot ventrad of seta D2 forms a whitish line extending anteriorly to 
the prothoracic shield. Prothoracic shield white with 6 black spots. Thoracic 
legs grayish-brown. Pinacula grayish-brown to brown. Lateral sclerites on ab- 

VoLUME 28, NUMBER 2 M57 

dominal prolegs light grayish-brown. Anal shield and lateral sclerites on anal 
prolegs dark grayish-brown. 

Fourth and Fifth Instar (Fig. 2 D). Head yellowish-brown suffused with 
light brown markings. Frons and adfrontals paler. Ocellar area black. Body 
creamy white with maroon transverse band on anterior half of meso- and meta- 
thorax and abdominal segments 1-9. Bands on mesothorax and 9th abdominal 
segment noticeably paler. Prothoracic shield whitish with 6 irregular black spots. 
Thoracic legs pale basally gradually darkening to grayish-brown on tarsus. Anal 
shield pale yellowish-brown. Spiracles dark brown with black peritreme. Pre- 
spiracular sclerite black, pinacula brown. 

Pupa (Fig. 2 A, B, C). Lightly sclerotized, light orangish-brown. Spiracles in 
shallow depression, rims of spiracles projecting above cuticular surface. Anterior 
margins of abdominal segments 5, 6, and 7 strongly pitted. Proboscis length variable, 
exposing metathoracic legs as figured or extending completely to apex of wings. 
Cremaster consisting of 2 elongate spines curving slightly ventrad. 


The assistance of Mrs. Sandra Shuler in various phases of this study 
and of Mrs. Phyllis Habeck for the illustrations is gratefully acknowl- 
edged. Identification of the wasp was by Dr. Eric Grissell and the spider 
by Dr. Karl Stone. 


Forses, W. T. M. 1954. Lepidoptera of New York and Neighboring States. Part 
3. Cornell Univ. Agr. Exp. Sta. Mem. 329. 433 p. 

Ganyarp, M. C. & U. E. Brapy. 1972. Interspecific attraction in Lepidoptera 
in the field. Ann. Entomol. Soc. Amer. 65: 1279-1282. 

Harpwick, D. F. 1958. Taxonomy, life history, and habits of the elliptoid-eyed 
species of Schinia (Lepidoptera: Noctuidae), with notes on the Heliothidinae. 
Can. Entomol. Suppl. 6: 116 p. 

KimsaL1, C. P. 1965. The Lepidoptera of Florida; An annotated checklist. Artho- 
pods of Florida and neighboring land areas. 1. Fla. Dept. Agr., Gainesville. 
363 p. 


A description of the larva of Isoparce cupressi (Bdv.) was given by the author 
(1973, J. Lepid. Soc. 27: 1-8), accompanied in the same issue by a field note 
by Van Buskirk (p. 83-84). Both articles omitted mention of a larval character 
brought to notice in the caterpillars collected by Van Buskirk near McClellanville, 
South Carolina. In my description of the development of the larva, much at- 
tention was given to the brown on the dorsal stripe and on the spiracular areas. 
In Van Buskirk’s specimens, many of them showed no such brown except for the 
dorsal horn and on the second thoracic spiracle. Instead, the areas mentioned showed 
the same yellowish white of the lateral lunules. All degrees between the two 
extremes were seen in the wild larvae. Van Buskirk’s wild caterpillars included 
various instars collected over a very few days, and so represented the offspring of 
several different females, leading to the conclusion that the natural coloration of 
the larva is variable in this respect. 

RicHarp B. Dominick, The Charleston Museum, Charleston, South Carolina 29401. 



Roperick R. Irwint 
24 East 99th Place, Chicago, Illinois 60628 

Of the ten butterfly names proposed or treated by Phélipe Poey in 
his Centurie de Lépidoptéres de Vile de Cuba (1832), three are today 
applied to species occurring in the Nearctic fauna: “Terias” [Eurema] 
dina Poey, “Eumenia” [Eumaeus] atala Poey, and “Melitea” [Phyciodes| 
frisia Poey. In only the last species is the nymotypical race Nearctic. 
Three other names, “Polyommatus” [Hemiargus hanno] filenus Poey, 
Pieris ilaire Godart [= Appias drusilla (Cramer)], and “Callidryas” 
[Phoebis] orbis Poey, have in the past been applied to North American 

There is general lack of agreement among catalogues of North Ameri- 
can butterflies in citing this work. Although Poey illustrated only a 
single species on each plate, in some references as many as three species 
are given the same plate number. These discrepancies are readily 
observed in Table 1, which contains exact quotations of the citations 
of the work by various catalogue compilers. 

Poey’s work was issued in parts, with unnumbered pages and plates, 
which were intended to be rearranged, numbered and bound on comple- 
tion of the work, but it was never finished; only two of the ten projected 
decades of ten species each were issued. The work was recently (“1970” 
[1971]) reprinted by E. W. Classey Ltd. Colonel Charles F. Cowan 
informed me (in litt.) that he has seen three original copies of Poey, 
the one that was reproduced by Classey and two others, and in all the 
order of species is the same. However, copies exist in which the species 
are arranged differently. Through the kindness of Ms. Carolyn Jakeman 
of the Houghton Library of Harvard University, such a copy was located 
in that library. The arrangement of species in this copy is shown in 
Majolke Il. 

Cowan (in litt.) suggested that the discrepancies noted may have 
resulted from the use of such a differing copy by an early author, pos- 
sibly Scudder, and the repetition, without checking, of his references by 
subsequent compilers. The rarity of Poey’s work may have prevented 
these normally careful authors from checking their references. The 
Classey text, agreeing in arrangement with most originals, will probably 
be accepted as definitive. 

1 Research Affiliate, Illinois Natural History Survey, and Honorary Curator of Lepidoptera, 
Illinois State Museum. 



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I gratefully acknowledge assistance provided by F. M. Brown, Lee 
D. Miller, and particularly Col. C. F. Cowan, whose ideas and sugges- 
tions are largely embodied in this note. 


Dyar, H. G. “1902” [1903]. A list of North American Lepidoptera and key to 
the literature of this order of insects. Bull. U.S. Nat. Mus. 52. 723 p. 

Epwarps, W. H. 1877. Catalogue of the Lepidoptera of America north of Mexico. 
Trans. Amer. Entomol. Soc. 6: 1-67. 

“1884” [1885]. Revised catalogue of the diurnal Lepidoptera of America 
north of Mexico. Trans. Amer. Entomol. Soc. 11: 245-337. 

Pory, P. 1832. Centurie de Lépidoptéres de Vile de Cuba, contenant Ja description 
et les figures coloriées de cent espéces de papillons nouveaux ou peu connus, 
représentés d’aprés nature, souvent avec la chenille, la chrysalide, et plusieurs 
détails microscopiques. J. A. Mercklein, Paris. (2 livraisons only, of 10 
projected.) 20 pls. 54 p. text (unnumbered). Reprinted “1970” [1971] by 
E. W. Classey Ltd. 

ScuppvER, S. H. 1875. Synonymic list of the butterflies of North America, north 
of Mexico. Part I. Nymphales. Bull. Buffalo Soc. Nat. Sci. 2: 233-269. 
1876. Synonymic list of the butterflies of North America, north of 

Mexico. Part II. Rurales. Bull. Buffalo Soc. Nat. Sci. 3: 98-129. 

SKINNER, H. 1898. A synonymic catalogue of the North American Rhopalocera. 
Amer. Entomol. Soc., Philadelphia. 100 p. 

STRECKER, H. 1878. Butterflies and Moths of North America. B. F. Owen, Reading, 
iPas2oon p: 


Megathymus yuccae yuccae (Boisduval & Le Conte) was described in 1833 from 
Aiken County, South Carolina. This skipper was known in South Carolina only from 
the type locality until March 1971, when I located a small colony along Hwy. 174 
south of Adams Run in Charleston County. In the spring of 1972 I found a very 
large colony at Edisto Island State Park in Charleston County. This area is 12 miles 
from the Adams Run colony. In 1973, three empty “tents” were found in a stand 
of yucca plants northeast of Mt. Pleasant along Hwy. 17 again in Charleston County. 
Dr. Douglas C. Ferguson captured a female on 4 April 1971 at the Wedge Plantation, 
Charleston County and Dr. Dominick collected two males there on the same date. 
In 1973 Dr. Dominick found a pupal tent in the same area, dug it up, acquired a 
live pupa and freeze dried it. The Edisto Island locality and the Wedge Plantation 
locality are at opposite ends of Charleston County, approximately 70 miles apart. 
The Mt. Pleasant area is approximately in the middle of the county. Thus Mega- 
thymus yuccae appears well established in coastal Charleston County. 

In February of 1973 I decided to visit Edisto Island and dig up pupae of 
M. yuccae. This was done in the middle of February, which was a month earlier 
than I had ever dug up M. yuccae before. (By February the larvae of M. yuccae have 
stopped feeding and tend to stay in the upper portions of the hostplant. A few 
days before the larvae pupate they stop moving in the typical “caterpillar” motion 

VoLUME 28, NUMBER 2 161 

and start propelling themselves up and down the tunnel in the cortex of the plant by 
rotating their abdomen, as is the character of the pupae.) I found several plants 
which had the tents of M. yuccae in them and started to dig. After I had dug up 
10 plants I decided to stop and come back at a later date to dig up the rest because 
I had found only 2 pupae, the rest being still in the larval state. Six of the 8 larvae 
were ready to pupate, the other two had discontinued feeding but were still active. 
On 17 and 18 March I again visited Edisto Island and dug up 25 more immatures. 
Of the 25 in the March batch only 3 were still larvae, the rest having already pupated. 
All 3 larvae were ready to pupate. 

I divided all of the immatures into two groups: group A (being those which were 
found in February); and group B (those found in March). Group A consisted of 
8 larvae and two pupae. Of the 8 larvae, 2 died in that state; the rest pupated but of 
these only two emerged as perfect adults, both females on 22 March. The other four 
emerged (3 males, 1 female) but the wings did not expand. Of the two pupae, one 
emerged a perfect male on 16 March. The other pupae had died when checked on 
16 March. 

Group B consisted of 22 pupae and 3 larvae. Of the three larvae, 2 had died 
when checked on 9 April. The other pupated, and a perfect female emerged on 
11 April. Of the 22 pupae, four died while 16 emerged as perfect adults (9 males 
on 21, 24, 26 (2), & 27 March and 8, 9 (2), 12 April; 7 females on 25, 29 March 
and 14, 15, 19, 20 and 21 April). Two others emerged but their wings did not 
expand (1 male, 1 female). 

The total sample of 11 larvae breaks down as follows: 4 died (36.5%) and 7 
pupated (63.5%). Of those which pupated, only 3 emerged as perfect adults 
(43% ) and 4 emerged deformed (57%). Of the 24 immatures which were taken in 
the pupal state: 5 died (21%), 17 emerged as perfect adults (71%), and 2 emerged 
deformed for 8%. In all, 17 of 24 pupae succeeded in reaching the perfect adult 
state (71% success). On the other hand only 3 of 11 larvae taken emerged as perfect 
adults (73% failure ). 

The conclusion is obvious, if you plan to collect Megathymus yuccae by digging up 
the immatures you will have far better success if you wait until the larvae have 
pupated. 71% success is better than 73% failure any day. It will be interesting to 
see if this mortality rate occurs in other species of Megathymidae as well. 


FREEMAN, H. A. Systematic Review of the Megathymidae. J. Lepid. Soc. 23, Suppl. 
1208 p: 

RONALD R. GATRELLE, 126 Wells Road, Hanahan, South Carolina 29405. 


When Mitoura hesseli (Rawson & Ziegler) was recognized as a new species from 
Lakehurst, New Jersey in 1950, several specimens from North Carolina were found 
by the late Frank Morton Jones in his collection. Since their capture in 1911, these 
had been assumed to be Mitoura gryneus (Hubner). These specimens (two males 
and two females) were captured on 28 July near Southern Pines in Moore Co. To 
my knowledge these are the only records from North Carolina. Therefore I was 
excited when I found M. hesseli at two locations in North Carolina during 22-25 
July 1972. The two locations, one on the Ft. Bragg Military Reservation and the 
other near the town of Raeford, are in Cumberland and Hoke counties respectively. 
Both of the counties border on Moore Co. 

I tried to find M. hesseli in April 1972 by visiting a number of the more accessible 


concentrations of White Cedar (Chamaecyparis thyoides) on the Ft. Bragg reserva- 
tion, but had no luck, and decided not to make an effort to look for the second 
brood. In July I was collecting hesperids that were visiting Sweet Pepperbush 
(Clethra alnifolia) when I spotted and captured the first M. hesseli also visiting these 
flowers. After realizing what I had captured, I made a quick search of the area and 
found a small stand of White Cedar (10-12 trees) about 20 yds. from the edge of a 
powerline cut in which I was collecting. More White Cedar may have been farther 
back in the wooded area. An hour’s worth of additional searching turned up 2 
more specimens. 

Having found my first M. hesseli I checked other promising stands of White 
Cedar and eventually found the second locality. The two areas in which M. hesseli 
were found were the only areas that had a considerable amount of Sweet Pepperbush 
in the vicinity of the White Cedar. I tried tapping trunks and throwing sticks 
into the upper branches of the White Cedar, but never saw M. hesseli on its foodplant. 
In both areas M. hesseli was uncommon, and a two hour search would turn up 
4-5 specimens. The patches of Sweet Pepperbush could be searched in 10—15 
minutes so that most of the time was spent just waiting for M. hesseli to appear on 
the flowers. Most collecting was done during midday, however, some specimens were 
taken as late as 1700, and almost all were in good condition. 

White Cedar is not uncommon along stream banks and in swamps in this part of 
North Carolina, and in view of the fact that the captures were made over a three 
county area separated by as much as 23 air miles, further collecting in this part 
of the state should tum up additional locations. Also it seems that the most promising 
areas to investigate, in July at least, would be those where Sweet Pepperbush or 
other productive flowers are in the vicinity of White Cedar. 

It is interesting to note similarities in the occurrence of M. hesseli in North 
Carolina and in New Jersey. Rawson & Ziegler (1950, J.N.Y. Entomol. Soc. 58: 
69-80) noted that M. hesseli in New Jersey was almost always found on flowers, 
and was not seen arriving or leaving, or flying about the foodplant as does M. 
gryneus. This certainly was the situation in North Carolina. 

The range of M. hesseli has now been extended to Virginia with the capture of a 
female on 18 July 1972 in Chesapeake. Mr. Bill Smith captured the specimen on 
milkweed (Asclepias syriaca), along with a female M. gryneus, while collecting on 
the eastern edge of the Dismal Swamp. 

There appears to be an unpublished record of M. hesseli from Maryland. In the 
correspondence of Mr. Frank Jones to Mr. J. B. Ziegler regarding the identification 
of M. hesseli in his collection, Jones cited a specimen from Pocomoke, Maryland, 
dated only 21 July and bearing a dos Passos identification label. The present status 
of the specimen or why the record has never been published before is unknown. 

M. hesseli may occur in many areas along the coastal region of the eastern U.S., 
but probably has remained uncollected not only because White Cedar often grows 
in inaccessible swamp lands, but also because M. hesseli is inconspicuous in areas 
where it does occur. 

Ricuarp A. ANpERSON, 1044-B Halsey Drive, Key West, Florida 33040. 


Interspecific hybrids between the Limenitis arthemis-astyanax complex and Limenitis 
archippus (Cramer) are extremely rare in nature. They occur as morphs preserving 
either more arthemis-like (arthechippus Scudder) or astyanax-like (rubidus Strecker) 
phenotypic characters. Each of these can be generally separated into two sub- 

VoLUME 28, NUMBER 2 163 

1 Cm: 

Fig. 1. Limenitis £. arthechippus from Stevens Point, Wisconsin. Right: right 
wings, dorsal; Left: right wings, ventral. Note that the photos are not exactly to 
same scale. 

morphs, light and dark. These interspecific hybrids have been reviewed by Platt, 
Frearson & Graves (1970, Can. Entomol. 102: 513-533) and Platt & Greenfield 
(1971, J. Lepid. Soc. 25: 278-284). 

An interspecific male hybrid of Limenitis, representing a ninth wild-caught Nearctic 
record of the hybrid morph arthechippus was captured by James M. Malick at Stevens 
Point (Portage County), Wisconsin, on 8 August 1961. It has been noted in a regional 
faunal study (Johnson & Malick 1972, Reports on the Flora and Fauna of Wisconsin 
7: 1-6) and deposited in the Museum of Natural History, University of Wisconsin, 
Stevens Point. Unfortunately, the genitalia of the specimen were accidentally 
destroyed after examination. The purpose of this paper is to report and describe 
the specimen, and speculate about what type of cross it represents. 

Platt & Brower (1968, Evol. 22: 699-718) and Platt, Frearson & Graves (op. cit.) 
have demonstrated that banded L. a. arthemis (Drury) and unbanded L. a. 
astyanax (Fabricius) are conspecific. Intergrades of this complex show continuous 
variation which may be divided into six categories. Categories 1 and 6 are the 
respective parental types; categories 3 and 4 are partially banded morphs referable 
to the form name proserpina Edwards; and categories 2 (banded) and 5 (un- 
banded) are more applicable to the name of each parent type. Genetic studies to 
date support the hypothesis that the white banding of arthemis is controlled by a 
major autosomal gene, the alleles of which display incomplete dominance (A. P. 
Platt, pers. comm.). Platt & Brower (op. cit.) have suggested that this complex 
exhibits primary, rather than secondary, intergradation, and that their populations 
in the “blend zones” are held in Hardy-Weinberg equilibrium by the neutralizing 
effects of selection favoring mimicry (astyanax) in southern regions and disruptive 


L R 

Fig. 2. Distal ends of the valvae, outer lateral view: L (left), R (right). The 
“hybrid” spines are indicated by the arrows. 

coloration (arthemis) northward. The intergrade forms presumably survive through 
some selective advantage of partial banding within the region where these selective 
forces become reversed (A. P. Platt, pers. comm. ). 

Random sampling of the genus at the approximate latitude of 44°32’ in Portage 
and Clark counties in Wisconsin indicates a ratio of 0.79 banded (arthemis)/0.08 
partially banded (proserpina)/0.13 unbanded (astyanax) for 24 specimens. This 
latitude is far enough north to expect the scarcity of the latter two morphs. A 
recent sampling of 100 specimens at Minneapolis, Minnesota (approx. latitude 45° ) 
(Bergman & Masters 1971, Mid-Continent Lepid. Ser. 2(31): 1-11) reflects a re- 
spective ratio of 0.58/0.20/0.22 for the three forms. L. archippus is commonly 
represented in samples taken from the exact collecting locality of the Stevens Point 

The Stevens Point hybrid (Fig. 1) is unusual because its coloration seems most 
like that of a proserpina intergrade. This was the tentative identification given it 
by Dr. Platt (from color slides) after its discovery. Only the subsequent genitalic 
dissection confirmed that the specimen is referable to arthechippus. Interestingly, the 
coloration of the wings is very similar to a group of proserpina noted as “form (c)” 
in an early review by Field (1904, Psyche 11: 1-6). Such specimens of proserpina 
are large, brown in ground color, have narrow and irregular mesial bands, and show 
prominent red spots on the surface of the secondaries. The new Wisconsin hybrid 
is large (wing-span = 5.6 cm) and its most distinctive character is the extremely dark 
ground color of the wings, like that of proserpina. The mahogany coloring so char- 
acteristic of other interspecific hybrids is reduced. 

Outline drawings of the distal ends of the valvae (Fig. 2) were traced from 
photos taken before the dissection was sent to Dr. Platt. He confirmed the dissection 
as arthechippus. Each valva clearly shows a distal downward pointing, blunted spine 
representing the intermediate condition between the short teeth of the arthemis- 
astyanax complex and the long downward curving and pointed spine characteristic 
of archippus (see Chermock 1950, Amer. Midl. Nat. 43: 513-569; Nakahara 1924, 
Bull. Brooklyn Entomol. Soc. 19: 166-180; and Platt, Frearson & Graves, op. cit., 
Fig. 3). This evidence confirms that the present specimen is an interspecific 
hybrid. The spine is bifurcate on the left valva, whereas, the right one is somewhat 
longer and narrower than that figured by Platt, Frearson & Graves (op. cit.). 
Further research on variability in the distal spines of these interspecific hybrids might 
link certain morphological characters with specific genotypes. 

VOLUME 28, NUMBER 2 165 

The Stevens Point specimen represents an aberrant morph, when compared to lab- 
reared or other wild-caught interspecific hybrids of Limenitis. The aberrant pheno- 
typic characters of this specimen suggest that it might represent a case of natural 
hybridization between an intergrade of the arthemis-astyanax complex and. archippus, 
or possibly the backcross of a male arthechippus to the arthemis parent (no females 
of arthechippus are known, presumably due to heterogametic inviability ). 

Two recent interspecific crosses of female proserpina intergrades with archippus 
males in the laboratory have produced a 1:1 ratio of arthechippus and rubidus siblings 
(A. P. Platt, pers. comm.). However, the above sampling ratios, and possibly the 
reduction of mahogany ground color in the wild hybrid, suggest that the latter 
speculation is more probable. 

I am greatly indebted to Dr. Austin P. Platt, University of Maryland Baltimore 
County, for suggestions, materials, verifications, and review of the manuscript. 
Thanks are also due to Dr. Charles A. Long, Director of the Museum of Natural 
History, University of Wisconsin, Stevens Point, and Mr. Peter L. Borgo, University 
of Delaware, for their consideration and comments. Father Roy Parker, Holy 
Cross, inked the drawings. 

Kurt Jounson, Museum of Natural History, University of Wisconsin, Stevens 
Point, Wisconsin 54481. Present address: (Br.) Novitiate, Order of the Holy Cross, 
West Park, New York 12493. 


Ten years of collecting moths in the Midwest has resulted in many interesting 
and unusual captures. Some of these species appear to be far out of their previously 
recorded ranges and these records may prove to be of interest to the researcher 
and collector alike. Among the most notable of these are Itame abruptata (Walker) 
a northern species previously known to occur in Canada and Northeast United States 
which was taken in Franklin Co., Missouri (5 and 7 June 1972), and in Washington 
Co., Missouri (6 June 1972) (several fresh specimens of both sexes); Euchlaena 
irraria (Barnes & McDunnough) another northem species before only known from 
Canada and as far south as Pennsylvania which was taken twice in Washington Co., 
Arkansas (27 May 1967 and 1972), and once in Franklin Co., Missouri (6 June 
1972) (all fresh males); Glena cribitaria (Guenee) an eastern species with the 
nearest previous record from its type locality of Northern Illinois was taken several 
times in Carroll Co., Arkansas (May 1965), Washington Co., Arkansas (April 1967, 
May 1966-9 and June 1971), Benton Co., Missouri (May 1970), and Washington 
Co., Missouri (June 1973) (in fresh series of males and females); Lytrosis sinuosa 
Rindge an eastern species with its nearest previous record from Oktibbeha Co., 
Mississippi was taken in Washington Co., Arkansas (4 June 1971, 27 May 1972) 
(fresh males), and Franklin and Washington Co., Missouri (5-7 June 1972) (in 
fresh series of males and females); Chloroteryx tepperaria (Hulst) a species of the 
Gulf States was taken in Washington Co., Arkansas (1 September 1968) (a single 
male), and 21 August 1971 (three fresh males); and Heterophleps refusata ( Walker) 
a northern and eastern species previously taken as far south as Virginia was taken 
twice in Clay Co., Missouri (29 May 1968, 4 May 1972), and once in DeKalb 
Co., Missouri (21 June 1972) (all fresh males). 

Rocer L. HerrzMan, 3112 Harris Avenue, Independence, Missouri 64052. 
(Research Associate, Florida State Collection of Arthropods, Division of Plant 
Industry, Florida Department of Agriculture and Consumer Services, Gainesville, 
Florida ) 



Based upon correspondence and conversations with collectors around the country, 
it would appear that many lepidopterists have only vague ideas concerning butterfly 
habitats in regions other than their home areas, unless they kave traveled widely. 
The purpose of this note is to point out that for a given species, extensive habitat 
variation may occur as a function of geography. To understand butterfly distribu- 
tions and subspeciation, collectors need to be aware of the interrelations among 
latitude, longitude, and altitude, as well as the more obvious factors such as annual 
mean temperature, rainfall and vegetation. A useful reference in these areas is 
Carpenter’s book (1956, An Ecological Glossary, Hafner, New York). 

To emphasize my point, several anecdotes and experiences are presented herein. 
It was once remarked to me that Laramie, Wyoming, my present home, with 
its elevation of 7100 ft. should be rich in arctic-alpine (tundra) species. When 
I asked why, the reply was that the top of Mt. Washington in New Hampshire is 
tundra at 6200 ft. and is populated by such species as Oeneis melissa semidea (Say) 
and Boloria titania montinus (Scudder). My eastern friend had neglected several 
factors. Laramie is a few degrees in latitude south of Mt. Washington and unlike 
New Hampshire, has an annual average rainfall of from 10 to 14 inches. The life 
zone is Upper Sonoran Desert or Bush-Steppe, Northwest Semi-Desert with mixed 
grasses and sagebrush as principal vegetation types. 

At Wyoming’s latitude (41° to 45° N), tundra or paramos occurs only above 
10,000 ft. As one travels north along the Rocky Mountains, tundra appears at lower 
elevations: about 6500-7000 ft. near Banff, Alberta (ca. 52° N) and 3000 ft. 
north of Fairbanks, Alaska (ca. 65° N). In this region of the Northwest, the 
longitude variation is from 105.5° W at Laramie to 145° W at Eagle Summit, north 
of Fairbanks. By contrast, along the northeast coast (64° to 72° W), tundra 
occurs on Mt. Washington (6288 ft., 71.25° W, 44.25° N), Mt. Katahdin, Maine 
(5268 ft., 68.9° W, 45.9° N), Mt. Albert, Gaspe, Quebec (ca. 4700 ft., 66° W, 
49.5° N). At Churchill, Manitoba, arctic tundra occurs at sea level (ca. 94° W, 
59o- N): 

The various subspecies of Oeneis melissa are tundra dwellers, whether they be 
found at high altitudes in the Rocky Mountains, or low altitudes in the Northeast 
and Far North. One cannot always make habitat projections of this nature, how- 
ever. In Maine and Quebec, and west through Minnesota, then North to the 
Northwest and Yukon Territories into Alaska, the subspecies of Oeneis jutta ( Hiibner ) 
are generally associated with muskeags (bogs), although in some areas of Alaska, 
jutta flies on the tundra above timberline. In Colorado and Wyoming, however, 
O. jutta reducta McDunnough is found in dry lodgepole pine (Pinus contorta var. 
latifolia Englem.) forests flying in the deadfall. Occasionally specimens are taken 
on flowers at the forest edge as reported by Ferris (1970, J. Lepid. Soc. 24: 306— 
307 ). 

An even more striking example of habitat variation occurs in Speyeria nokomis 
(Edwards). In the United States, nokomis is associated with very wet areas, either 
sloughs or moist alpine meadows where the larval foodplant, Viola spp., grows 
in the understory (Ellis 1969, J. Lepid. Soc. 23: 62-26; Ferris & Fisher 1971, J. 
Lepid. Soc. 25: 44-52). In Mexico, S. nokomis coerulescens (Holland) is found 
in a completely different habitat. The butterfly flies in Chihuahua and Durango 
in dry pine woods. During its flight season (late August and September), frequent 
rains maintain an almost bog-like condition of the pine needle and duff layer 
on the forest floor. Violets grow in this layer (L. P. Grey and J. R. Mori, pers. 
comm.). Adult nokomis are found in some nearby meadows, but only because of 
a concentration of thistles and other nectar sources. 

Latitude can play a major role in determining where butterflies of a given species 
are found. Two of many possible examples are: Papilio i. indra Reakirt and Lycaena 

VoLUME 28, NUMBER 2 167 

cupreus snowi (Edwards). The former butterfly is normally associated with ridge 
tops in foothill areas. This is certainly the case in the eastern Sierra Nevada and 
in Colorado and Wyoming. Occasionally specimens are taken at lower elevations 
at puddles following a rain shower. My initial experiences with indra and other 
subspecies confirmed that they occupy the barren ridge habitat where certain of 
the larval foodplants, Umbellifereae, grow. During the summer of 1972, I found 
that i. indra is a moist meadow flier in the Sawtooth Mountains of Idaho. Specimens 
were taken flying and nectaring in Transition Zone meadows at 7100 ft. The Idaho 
locality was about two degrees further north in latitude than the most northern 
point in Wyoming where I have taken indra. 

Lycaena cupreus snowi is recorded in Colorado only from above timberline (ca. 
10,000 ft. and above) where it flies next to the snow fields (Brown et al. 1957, 
Colorado Butterflies, Denver, Colo.). Just to the north, in Wyoming, the butterfly 
is generally taken in the Upper Transition and Canadian Zones in open meadows. 
I have taken a few specimens at timberline in the Wind River Mountains and on 
the Beartooth Plateau, but most specimens have been taken at much lower eleva- 
tions. In the Sawtooth Range in Idaho, I have found snowi relatively common in 
open meadows at 7100 ft. 

In conclusion, I would make three points. First, different subspecies of a butter- 
fly may be found in widely disparate habitats, in different parts of North America. 
Second, based upon observations from one locality, one cannot make inferences 
concerning other regions without a thorough knowledge of all of the environmental 
variables involved. Third, in some instances, “micro-environments” may be the 
dominant factor controlling species distributions, as in the Mexican race of Speyeria 

(This note is published with the approval of the Director, Wyoming Agricultural 
Experiment Station, as Journal Article JA 595.) 

CiirrorpD D. Ferris, College of Engineering, University of Wyoming, Laramie, 
Wyoming 82071. (Research Associate, Allyn Museum of Entomology, Sarasota, 
Florida; Museum Associate, Los Angeles County Museum of Natural History, Los 
Angeles, California ) 


I would like to make a few additions to J. A. Downes’ paper under this title 
(1973, J. Lepid. Soc. 27: 88-99). I discussed this question on more than one 
occasion with Collenette, and he often said that he was of the opinion that the 
attraction of damp mud lay in the mineral salts in general, and probably in the 
sodium chloride in particular, contained therein. I have two pieces of evidence 
that would appear to support this opinion. When collecting at Lech-am-Arlberg 
in Austria in 1958, it was noticed that, whilst the damp mud surrounding puddles 
formed in the mountain paths after a storm were highly attractive to butterflies, the 
damp mud at the side of permanent trickles of water crossing the same paths were 
always completely deserted. Presumably the mud round the puddles would be 
heavily impregnated with mineral salts leached out of the surrounding soil, whilst 
mud beside permanent trickles would have had everything washed out. Again in 
Africa it is noticeable that the attraction of damp mud increases with the distance 
from the sea. In the Shimba Hills I have only found Anthene lasti Sm. & Kby. 
(Lycaenidae) at damp mud; the nymphalids that feed at damp mud are the same 
species that are attracted to fermented fruit baits and, as both sexes are present, 
the attraction is obviously juices from rotten fruit that has fallen from the trees 
bordering the path. In the actual coastal forest at Jadini, situated within a mile 


from the sea, and in the Arabuku-Sekoke Forest, males of the various Papilio and 
Graphium species (Papilionidae) and Appias sabina Feld. (Pieridae) occur at damp 
mud, but nothing else. Both the latter forests are far more sheltered and _ less 
windy than the forests of the Shimba Hills. Compare this very meagre list 
with the number of species recorded in the Kenya Highlands and Western Uganda 
when motoring through in 1960: Kenya Highlands (Danaidae 3, Acraeidae 3, 
Nymphalidae 2, Lycaenidae 11, Pieridae 5, Papilionidae 1); Western Uganda 
(Danaidae 5, Acraeidae 0, Nymphalidae 5, Lycaenidae 7, Pieridae 9, Papilionidae 
3). (A full list of species will be found in 1962, Entomologist 95: 17-18). It is, 
perhaps, noteworthy that not a single hesperiid was recorded, as this is the family 
concerned in all the records of butterflies settling on human skin, exuding a drop 
of fluid from the anus, and then sucking it up through the proboscis. 

It is suggested that butterflies in the coastal areas absorb enough salt with their 
food as larvae, the shore level forests being more sheltered, and so receiving less 
salt, than the more exposed forests in the Shimba Hills. The air all along the 
Kenya coast is heavily impregnated with salt, which is deposited as a thin film 
on windows and which causes heavy corrosion in metal fittings. 

I was surprised to read in Downes’ paper that horse droppings were found to 
increase the attractiveness of damp mud and water. In Africa the droppings of 
herbivorous animals, antelopes, buffalo and elephant for example, hold no at- 
traction whatever, and it is only the droppings of carnivores that are attractive. 
I am not even certain that the faeces of the Canidae are attractive, as the droppings 
of my own dogs, which are fed mainly on meat, do not attract the Charaxes species 
that frequent my garden, but I have never come across the droppings of jackals 
or foxes in the bush. 

That there is some difference in the food requirements of male and female 
butterflies appears to be confirmed by the habits of the Charaxidae, where 
both sexes are attracted to fermenting fruit and sap but only the males to dung and 
carrion. Possibly there is some connection with the production of the female- 
attracting scents secreted by the males. A very specialised case of this connection 
is the recently discovered necessity for the males of certain Danaidae to feed on 
the fermented juices of certain plants of the Boraginaceae before they can develop 
this scent. 

I have one example of male moths feeding at damp mud, a Semiothisa sp. 
(Geometridae) identified by the British Museum (Natural History) as near 
fuscataria Mschl., which was found in considerable numbers at mud puddles in 
a forest in Uganda. 

D. G. SEvAsToPpuLo, F.R.E.S., P.O. Box 95026, Mombasa, Kenya. 


A great deal of information is available concerning the larval foodplants and 
parasites of the more important lepidopterous pests in Arkansas. Little such in- 
formation is available for Lepidoptera species not considered primary insect pests. 
With greater emphasis on pest management programs in recent years and increasing 
interest in biological control of pests, it is becoming more important to understand 
the relationships among animals and plants. In order to elucidate some of these 
relationships, the following study was conducted on or near the White River 
National Wildlife Refuge during the summer months of 1969 and 1970. 

The White River Refuge, a relatively undisturbed habitat, consists of portions 

VOLUME 28, NUMBER 2 169 

of Arkansas, Phillips and Monroe counties. The Refuge is 56 miles long, 3 to 6 
miles wide, and consists of 113,600 acres of mixed hardwoods. About 75% _ of 
the Refuge is flooded in winter and spring by the Mississippi, Arkansas, and White 
Rivers. Approximately 600 to 1000 acres are annually planted in agricultural crops, 
a portion of which serves as food for the wildlife. The crops include rice, soybeans, 
grain sorghum, other small grains and forage grasses. 

Lepidopterous larvae were collected on or adjacent to the Refuge and brought 
to a field laboratory for rearing. Each larva and a portion of the hostplant were 
placed in a waxed paper cup covered with a clear plastic lid. If the larva was 
an internal feeder, i.e. stem borer or leaf miner, the hostplant was placed in a 
vial of water within the cup. Fresh cuttings of the hostplant were added daily 
to the cups containing externally feeding larvae. The older food was removed 
only after the larvae had willfully changed to the new food. The cups were checked 
daily for emergence of adult moths and parasites. Hostplant specimens were col- 
lected for identification. 

Twenty-six lepidopterous species representing 12 families were reared from 
20 species of host plants. Seventeen species of parasites representing 4 families 
of Hymenoptera and Diptera were reared from the Lepidoptera. The following 
is a list of the lepidopterous species reared, dates of emergence, localities collected, 
hostplants and parasites. The dates given are dates of emergence of the moths and 
parasites. ; 

Acrobasis sp. PHYCITIDAE. 11 August 1970, 4 miles east Ethel. Host: Liquid- 
amber styraciflua (sweet gum). Parasites: Campoletis sp., Exochus sp., Trathala 
tetralophae (Cush.), ICHNEUMONIDAE;; Phanerotoma tibialis (Hald.), BRACONI- 
DAE; Pseudochaeta clurina Rein., TACHINIDAE. 

Anavitrinella pampinaria (Guenee) GEOMETRIDAE. 26 July 1971, 4 miles 
southeast Ethel. Host: Eupatorium sp.; 10 July 1971, 8 miles southeast Ethel. 
Host: Amaranthus hybridus (pigweed). 

Argyrotaenia velutinana (Walker) TORTRICIDAE. 4 July 1971, 5 miles south- 
east Ethel. Host: Ilex decidua (deciduous holly). 

Asterocampa clyton ( Boisduval & LeConte) NYMPHALIDAE. 12 September 1971, 
4 miles southeast Ethel. Host: Celtis mississippiensis (Hackberry). Parasites: 
Apanteles sp. BRACONIDAE. 

Brachmia melantherella (Busck) GELECHIIDAE. 17 July 1970, 2 miles south- 
east Ethel. Host: Xanthium pennsylvanicum (cocklebur). 

Characoma nilotica (Rogenh.) NOCTUIDAE. 12 July 1971, 3 miles north St. 
Charles. Host: Salix sp. (willow). Parasites: Brachymeria ovata (Say), Spilo- 
chalcis sanguineiventris (Cress.) CHALCIDIDAE. 

Choristoneura rosaceana (Harris) TORTRICIDAE. 30 June 1971, 5 miles south- 
east Ethel. Host: Hypericum sp. (St. John’s wort). 30 June 1971, 5 miles south- 
east Ethel. Host: Rubus sp. (blackberry). 

Conchylodes platinalis (Guenee) PYRAUSTIDAE. 27 July 1971, 5 miles north- 
west Snow Lake. Host: Ambrosia trifida (giant ragweed). 

Delta ramulosa (Guenee) NOCTUIDAE. 2 August 1971, 4 miles southeast Ethel. 
Host: Hypericum sp. (St. John’s wort). 

Desmia funeralis (Huebner) PYRAUSTIDAE. 13 July 1971, 5 miles southeast 
Ethel. Host: Vitis sp. (wild grape). Parasites: Eucordyligaster septentrionalis 

Eumarozia malachitana (Zeller) OLETHREUTIDAE. 19-23 August 1971, 3 miles 
southeast Ethel. Host: Diospyros virginana (persimmon). Parasites: Agathis 
annulipes (Cress.) BRACONIDAE. 

Fascista cercerisella (Chambers) GELECHIIDAE. 25 July 1971, 4 miles south- 
east Ethel. Host: Cercis canadensis (redbud). 

Filatima serotinella (Busck) GELECHIIDAE. 23 July 1971, 2 miles southeast 
Ethel. Host: Prunus sp. (wild cherry). 


Heterocampa subrotata (Harvey) NOTODONTIDAE. 3 July 1971, 5 miles south- 
east Ethel. Host: Celtis mississippiensis (hackberry ). 

Hymenia perspectalia (Huebner) PYRAUSTIDAE. 28 August 1971, 2 miles south- 
east Ethel. Host: Amaranthus hybridus (pigweed). 

Hysoropha hormos (Huebner) NOCTUIDAE. 19 August 1970, 5 miles southeast 
Ethel. Host: Diospyros virginiana (persimmon). Parasites: Meteorus sp. 

Loxostege sp. PYRAUSTIDAE. 27, 28 August 1971, 2 miles southeast Ethel. 
Host: Amaranthus hybridus (pigweed). Parasites: Cremnops haematodes ( Brulle) 
BRACONIDAE; Nemorilla pyste (Walker) TACHINIDAE. 

Mineola indigenella (Zeller) PHYCITIDAE. 1-3 August 1971, 4 miles southeast 
Ethel. Host: Crataegus viridis. Parasites: Eusisyropa boarmiae (Coq.), Eusisyropa 
virillis (Aldrich & Webber) TACHINIDAE. 

Olene leucophaea (Abbot & Smith) LIPARIDAE. 2 October 1970, 6 miles south 
St. Charles. Host: Liquidamber styraciflua (sweet gum). 

Phaecasiophora niveiguttana (Grote) OLETHREUTIDAE. 1 August 1971, 2 miles 
southeast Ethel. Host: Sassafras sp. (sassafras). Parasites: Macrocentrus ancy- 
livorus Roh. BRACONIDAE. 

Polychrosis sp. OLETHREUTIDAE. 4 August 1971, 2 miles north St. Charles. 
Host: Amaranthus hybridus (pigweed). Parasites: Agathis annulipes (Cress. ) 

Psilocorsis caryae Clarke OECOPHORIDAE. 23 August 1970, 3 miles east Ethel. 
Host: Carya sp. (hickory). 

Psilocorsis quercicella (Clemens) OECOPHORIDAE. 4 August 1971, 4 miles 
southeast Ethel. Host: Quercus sp. (oak). Parasites: Temelucha grapholithae 

Scythris trivinctella (Zeller) SCYTHRIDAE. 2 October 1970, 3 miles southeast 
Ethel. Host: Amaranthus hybridus (pigweed). Parasites: Nemorilla pyste 

Stegasta bosqueella (Chambers) GELECHIIDAE. 18-24 July 1971, 5 miles 
southeast Ethel. Host: Cassia fasiculata. 

Xenolechia “Telphusa” sp. group GELECHIIDAE. 2 August 1969, 2 miles south 
St. Charles. Host: Salix sp. (willow). 

Thanks are due Mr. Raymond McMasters for his cooperation in allowing use of 
the White River National Wildlife Refuge; Drs. Ed Smith and Patricia Coons for 
plant determinations; Drs. Paul Marsh, R. W. Carlson, C. W. Sabrosky, and B. D. 
Burks for determination of parasites; and Drs. R. W. Hodges and Ed Todd for 
determination of Lepidoptera. This article is published with the approval of the 
Director, Arkansas Agricultural Experiment Station. 

RicHarp L. BRowNn AND RopertT T. ALLEN, Entomology Department, University 
of Arkansas, Fayetteville, Arkansas 72701. 


At the request of Dr. Richard B. Dominick this small amount of information 
is offered to anyone interested in freeze-drying caterpillars without purchasing any 
materials whatsoever. In the summer of 1971 several saturniid caterpillars (mainly 
Hyalophora species and hybrids) were frozen alive with the original intention of 
keeping a small larval collection in the freezer permanently. On adding more speci- 
mens to the box in the spring of 1973, the 1971 ones were observed to be very 
light in weight. They were taken out, and no changes have been observed for 

VoLUME 28, NUMBER 2 7 

months thereafter. Some wrinkling of the skin had occurred in the freezer, but 
the colors were excellent, including tubercles. The fact that the kitchen freezer 
used was self-defrosting apparently answers the question of where the moisture 
went. It seems that had a well-ventilated box been used, the same results could 
have been achieved in several months instead of two years. 

RicHARrD S. PEIGLER, 303 Shannon Drive, Greenville, South Carolina 29607. 


Some time ago I reported on the process of freeze-drying and vacuum dehydra- 
tion for the preservation of immatures (Dominick 1972, J. Lepid. Soc. 26: 69-79). 
Since then, experience has led to some modification of procedure and equipment. 
These recommendations form the substance of this article. It is assumed that the 
reader has before him the previous report (of which a few reprints are still avail- 
able), for this article will proceed point by point on that basis. 

The pump oil should be changed regularly, otherwise the efficiency of the pump 
may be seriously impaired. An oil change is recommended after every 20 hrs. of 
operation, so a disconnect coupling (an “O” type ring) is desirable. Such disconnect 
couplings will also be found useful for anyone desiring to construct a mobile field 

The inside diameter of the tubing is of no great consequence in the system 
described. However, % in. tubing is recommended over % in., for two minor 
reasons: first, a slightly more efficient pull-down time will result, and second, 
the larger diameter is a bit easier for the amateur to flare or solder. 

Next, it is advantageous to lower the temperature of the freezer below the 
-7° C (20° F) previously recommended, since opening the freezer door can quite 
easily raise the temperature to above freezing. By removing the taped end of the 
thermostat from the ice-making compartment and gently bending it out of the 
way into the rear of the larger compartment, the temperature of the whole unit may 
be lowered to between —12 to —15° C (10 to 5° F), while the ice-making compart- 
ment goes down to about —24° C (-15° F). The resulting increase in drying time 
is not sufficient to be of practical concern. This lower temperature, in fact, is 
theoretically more suitable for the preservation of integrity of the cells. 

Previously the suggestion was made that Duco or similar cement would help 
preserve the integrity of the permanent joints. The suggestion is erroneous, for 
proper flaring alone guarantees the adequacy of sealing. If the flare (or soldering) 
is not properly made, no amount of posthumous treatment will help. 

As for killing the animal, I have largely abandoned the method of very quick 
deep freezing, which often agitates the larva so much that presentation of a lifelike 
attitude becomes difficult. It also influences the cellular integrity by destructive 
crystal formation. Slow freezing in the main compartment in general seems best. 
In case an undesirable attitude prevails, correction should be made as soon as 
possible, before the larva is frozen through. Try to avoid thawing a frozen specimen, 
for this has undesirable effects on some of the color pigments. Try to manipulate 
the larva when it is just cold enough to be dormant, but before cellular freezing. 
Frank R. Hedges, Houston, Texas, suggests that contact with the ambient air after 
any degree of freezing might change some of the chemically activated color pigments, 
and my own experience tends to bear this out. In such a case the larva may be 
put straight away into a cold desiccator and left to freeze to death, applying vacuum 
only when thoroughly frozen. One may have to sacrifice a lifelike posture in favor 
of coloration. More experimentation is needed. Other methods of killing are 


satisfactory, for example, cyanide or boiling water. With regard to ethyl acetate 
and other organic solvents, one must consider the possibility of solubilizing 
effects on certain plastics used in the apparatus. Properly processed larvae tend 
in general to retain their color well, with the exception, in my limited experience, 
of some greens and a few reds. 

Concerning the equipment, first there are the desiccators. I now use exclusively 
the Nalgene vacuum chamber with neoprene gasket. It is made of transparent 
polycarbonate and may be ordered with gasket and plastic top. It stands about 
25 cm high, and the recommended freezer holds two with ease. A further blessing 
is that they need not (in fact must not) be greased. If one is careful to keep the 
contact surfaces free of dust, dirt and ice crystals, these desiccators will hold the 
vacuum very well. A further advantage is that the plastic will not break and 
splinter to the dangerous proportion of a glass vessel in case of an implosion. 

As for pumps, valves and manometers as well as all the fittings, I have recently 
been in touch with a company whose catalogue offers such equipment of commend- 
able quality at good prices. For example, I have been told by two refrigeration 
experts that the pump I now use is rated at a vacuum of 0.1 micron, but will not 
pull down in practice to better than 25 microns/Hg, which is still adequate for the 
purpose. To understand the need for a high vacuum efficiency in the process, one 
must realize that 1 micron equals approximately 1/25,400 in. Hg, and remember that 
any pressure abcve 1-200 microns in the system renders the vacuum operation 

Some companies also supply high vacuum line valves, copper tubing, “O” type 
disconnect fittings, and for the permanent joints, a method of soldering requiring 
only the heat from a small propane torch. Such a method of fixing the permanent 
joints, of course, eliminates the task of flaring, a job extremely difficult to accomplish 
successfully where there are numerous joints in close proximity to one another. 

Practical suggestions as to specific companies have been published recently in 
the News. 

My warm thanks to Dr. Hermann A. Flaschka, who has taken time to edit and 
correct the original manuscript with humor as well as detail, and to Dr. Theodore 
D. Sargent, who has performed further needed surgery. 


RicHARD B. Dominick, The Charleston Museum, Charleston, South Carolina 29401, 


Saturnia walterorum Hogue & Johnson is perhaps the rarest saturniid in the United 
States, occurring locally in southern California. There are relatively few field data 
available for this moth. Fewer than 30 specimens have been collected and most 
specimens in collections have been reared from eggs secured from captured females. 
Sala & Hogue (1958, Lepid. News 12: 17-25) described the life history of S. 
walterorum reared under laboratory conditions. It is the purpose of this paper 
to present new information on flight period, distribution, and larval host records. 
A future publication will examine the taxonomic relationship between S. walterorum 
and S. mendocino Behrens. 

Saturnia walterorum is known only from 4 coastal counties in southern California: 
San Luis Obispo, Los Angeles, Orange, and San Diego. One specimen in the 
Los Angeles County Museum of Natural History is labeled “Cajon Valley.” This 
label may refer either to El Cajon Valley in San Diego County or Cajon Pass in 
San Bernardino County. Suitable habitats seem to exist in at least 4 additional 

VOLUME 28, NUMBER 2 173 

Fig. 1. Known distribution of Saturnia walterorum in southern California. 

counties in southern California and portions of Baja California, but no available 
records indicate that it has been sighted or collected other than in the previously 
mentioned counties. The moth has been collected at elevations varying from 100 
to 5500 ft. on both the coastal and high desert slopes of the mountains in southern 
California (Fig. 1). The flight period at lower elevations begins in late February 
and extends into mid-April, while at higher elevations specimens have been 
collected between April and mid-May. Individuals are on the wing only on warm 
sunny days from 0930 to 1530 hrs. 

The larval hostplant of S. walterorum has been in question for some time. Field 
observations indicate that larvae, pupae, and oviposition have occurred only on 
plants of the families Anacardaceae and Ericaceae. It is interesting to note that 
plants of the family Ericaceae serve as the host for S. mendocino, a species which 
occurs to the north of, and is thought to be allopatric to, S. walterorum. 

Two species of Anacardaceae are known to serve as natural larval hosts; these are 
Rhus laurina Nuttall and R. integrifolia Bentham & Hooker. Both species of Rhus 
inhabit dry chaparral slopes below 3000 ft. On two occasions females were observed 
ovipositing on R. integrifolia by the author, and empty cocoons have been found 
in association with this shrub (Sala & Hogue, op. cit.). Larvae have also been 
collected on R. lauwrina in San Diego by R. Hatch (pers. comm.). 

At higher elevations it appears that members of the family Ericaceae serve 
as larval hosts. One larva has been collected on an unidentified species of 
Arctostaphylus in the San Gabriel Mountains (Sala & Hogue, op. cit.). Adults have 
been observed in association with various species of Arctostaphylus in the Santa 


Monica, and San Gabriel Mountains by C. Henne (pers. comm.) and in the 
Laguna Mountains by R. Breedlove (pers. comm.). 

The larvae of S. walterorum exhibit two distinct color phases; one green, and 
the other reddish-orange. The bark of Arctostaphylus and areas of new growth on 
both species of Rhus are characteristically red in color. Therefore, each color phase 
of the larvae may blend into different portions of its environment. This adaption 
may make them less conspicuous to natural enemies while on the hostplant. Larvae of 
both color phases emerged from the ova of one female in a ratio of 1:1. 

The cocoon, which is brown and coarsely constructed, appears similar to the 
dried inflorescences of Rhus. This similarity may be advantageous, for reared 
larvae usually pupate at the terminal ends of branches among dried flowers or at 
the base of the plant, thus concealing the pupation site. 

I would like to thank the following individuals for allowing me to examine their 
records and specimens: Christopher Henne of Pearblossom; Fred Thome of E] Cajon; 
Dave and Jean Roldness of San Diego; James Tilden of San Jose; Eric Walters of 
Anaheim; and Charles Hogue and Julian Donahue of the Los Angeles County 
Museum of Natural History. 

P. M. Tuskes, Department of Entomology, University of California, Davis, Cali- 
fornia 95616. 


For a number of years my sons and I have been breeding butterflies found in 
El] Salvador. Among them we have reared from egg to adult, several times, groups 
of Battus polydamas polydamas L. The full process has taken an average of 40 
days, pupation alone from ten to eighteen days. According to Young (1971, Ann. 
Entomol. Soc. Amer. 60: 595-599), in Costa Rica the total development for 
this species averages 41.32 days, ranging from 36 to 46 days, with pupation ranging 
from 14 to 16 days. 

On 14 June 1972 we saw a female lay seven eggs, which were collected and 
put in a transparent plastic bag. On the 20th, the eggs hatched. The larvae were 
fed with fresh leaves of the foodplant, Aristolochia anguicida L., until pupation, 
which occurred between 8 and 10 July. Only five larvae had survived out of 
the seven. On 17 July, the first adult, a male, emerged, that is 9 days after pupa- 
tion, which is a little shorter than usual. The next adult, another male, emerged 
on 19 July, 11 days after pupation, being this time closer to average. 

With the remaining three individuals, pupation time was completely unexpected. 
The third adult, a male again, emerged on 27 October; the fourth, another male, 
on 7 December; and the fifth, a female, on 25 January 1973! The pupal stage 
in these three cases was 110, 150 and 199 days respectively. All of these adults 
were absolutely normal and healthy. 

We emphasize the fact that this species is gregarious during the early stages, and 
that the seven larvae were kept in the same bag, and therefore under the same 
conditions of food, light and temperature. When pupation occurred, the pupae 
were placed in the same pupation box, and again they were exposed to the same 
environmental conditions until the first two adults emerged. At that time one 
of the pupae was given to Mr. Steve R. Steinhauser, who lives in the neighboring 
town of Santa Tecla, some 13 km. from San Salvador, at a slightly higher altitude. 
This pupa was the one that lasted 199 days. The other two were in the same box 
at all times. 

VoLUME 28, NUMBER 2 175 

We have found a report of aestivation in one species of Saturniidae, Rothschildia 
lebeaui ? aroma Schaus (Quezada 1967, Rey. Biol. Trop. 19: 211-240), whose 
pupa spends the six months of the dry season waiting for the first rains to fall. The 
dry season in this country starts in November and ends in April. The wet season 
starts in May and ends in October. Consequently the case we are reporting hap- 
pened during part of the wet season and part of the dry. It is true that the 
weather was somewhat chaotic during 1972, there being a long spell of dry weather 
during July (20 days) and August (15 days), and then copious rain during 
November and the beginning of December. 

ALBERTO MuysHonpt, 101 Avenida Norte 322, Lomas Verdes, San Salvador, El 


My notes for the Spring of 1973 show that cardui first came to my attention at 
Fountain Valley School (11 miles SE of Colorado Springs, Colo.) on 19 April. That 
day had started with near freezing temperatures and a light snow flurry. At noon 
I saw two cardui flutter over the lawn. A week or ten days of warmish weather fol- 
lowed. with early morning temperatures as high as 44° F. Around 1430 on 28 April 
I was driving south from Denver on State Highway 83. About five miles north 
of Parker I met swarms of cardui drifting toward the northeast. I estimated about 
100/150 passing directly in front of me each mile. This continued almost all 
of the way home, about 70 miles. There was a break going over the Platte-Arkansas 

Examination of the insect, as flying at Fountain Valley, showed the specimens 
to be badly worn to tattered and the ground color to be quite pale. The con- 
centration on the lawns may have reached 500 per acre. These butterflies were 
feeding at dandelion flowers and later apple blossoms. The numbers held reasonably 
steady until 11 May when another wave arrived. These specimens were consider- 
ably larger, and much fresher and darker in color. A careful estimate made around 
1530 that afternoon placed the numbers on the lawns (30 acres) at about 1,000 
per acre and on the prairie (2,000 acres sampled) about 150 per acre. These 
concentrations remained relatively constant until the weekend of 19-20 May and 
the two succeeding days when the weather was rainy and cold. By 23 May there 
were very few cardui around, just about the normal situation. 

On 18 May driving south on I-25 to New Mexico, the numbers of cardui flying 
across the highway were high enough to materially reduce the efficiency of the 
automobile’s radiator. It was necessary to stop after about 100 miles to clean 
the radiator and scrape the squashed remains from the windshield. This situation 
continued through the 18th and 19th. 

There are very few thistles in the vicinity of Fountain Valley School. It will 
interesting to see if we have an abnormally large crop of cardui in early summer. 
If we do, it will be important to discover the alternate foodplant here. The several 
large patches of thistle known to me along highway 83 in Douglas County will be 
watched with interest. 

Postscript: Larvae used Helianthella and two species of Lupinus after the few 
thistles were stripped to the ground. 

F. Martin Brown, Fountain Valley Rural Station, Colorado Springs, Colorado 



On 24 May 1973 near McClellanville, South Carolina, an Anisota virginiensis 
pellucida (J. E. Smith) female emerged from a brood reared on Quercus nigra 
the previous August. She was put outdoors to call in males during the day. Other 
matters pressing, I was able to spend only short periods at the cage, during which 
time I saw several males of the same species and also captured three males of 
Amphion nessus (Cramer) that homed in directly to the cage and buzzed about 
trying to get in. The data follow, all times being Eastern Standard time: 

Anisota virginiensis 6 6: 24 May, 1045; 25 May, 1107 and 1130. 

Amphion nessus 6 6: 26 May, 1415, 1416 and 1430. 

The most obvious possibility would seem to be a similarity in the chemical con- 
figuration of the sex pheromones of the two species, distantly related as they are. 
(A previous paper (Dominick, R. B. & C. R. Edwards 1971, J. Lepid. Soc. 25: 
84-85) reported on the flight pattern of male Anisota virginiensis. ) 

RicHArD B. Dominick, The Charleston Museum, Charleston, South Carolina 29401, 


THE EvoLurion oF MELANISM, The Study of a Recurring Necessity, With Special 
Reference to Industrial Melanism in the Lepidoptera, by Bernard Kettlewell. 1973. 
Clarendon Press, Oxford. xxiv + 424 p., illus. + plates. Price: $33.00 (U.S.). 

This eagerly anticipated work provides a valuable compilation of the data and 
conclusions of Kettlewell and his associates on the phenomenon of melanism in the 
Lepidoptera. Although broad in scope and rich in detail, the book possesses some 
shortcomings which will be discussed following a résumé. 

The work is divided into 19 chapters (in seven major sections), followed by 
three appendices, a list of recorders, and a bibliography with better than 600 
entries (including references through 1971). There are 38 pages of plates (35 
halftone, 3 color), 14 text figures, and some 40 tables. 

The book begins with a general consideration of melanism, its nature and func- 
tions (3 chapters). This introduction stresses Kettlewell’s major theme that melanism 
has been a recurring necessity in the evolutionary histories of diverse organisms, This 
section is followed by one on melanism specifically in the Lepidoptera (3 chapters), 
which includes classifications of both adult and larval melanisms, as well as a general 
treatment of the phenomenon of industrial melanism, and a review of the world- 
wide distribution of that phenomenon. 

Attention is then focused on the now-famous Biston betularia (3 chapters). The 
mark-release-recapture selection experiments in Birmingham and Dorset (1953- 
1955) are recounted, and the history and spread of the melanic forms in Great 
Britain are documented. Special reference is made to the frequency surveys (1952— 
1970) which Kettlewell has compiled from the records of nearly 170 observers, 
and these data are detailed in an 11l-page appendix. Kettlewell then turns to 
consider non-industrial melanisms (3 chapters), in particular his own extensive 
work, including mark-release-recapture experiments, on Amathes glareosa in Shet- 


land. This section is followed by a treatment of recessive melanism (2 chapters ) 
in which recent work on Lasiocampa quercus is described. A variety of melanisms 
are then described as miscellaneous (4 chapters). This last section includes examples 
of aposematic, sex-linked, and environmental melanism, as well as a short considera- 
tion of melanism in butterflies. The main body of the text is concluded with a 
regrettably short synthesis (1 chapter, 6 p.), and there follow appendices on 
breeding techniques (4 p.) and melanism in British moths (38 p.). 

To turn now to criticisms of the book, I would first point out certain matters 
which may provide some annoyance to readers. The most important of these 
concerns the arrangement and numbering of the “plates.” These plates (actually 
halftone figures) are numbered in the order of their citation within the text, but 
are arranged into bundles of halftone pages in an oftimes different sequence. Thus, 
one finds for example, plates 3.1 and 7.2 on the same halftone page. I sought plate 
5.17a for a full five minutes after coming to its citation in Chapter 5. (The situa- 
tion is rendered more confusing in my copy of the book by an error involving 
reversal of the plates belonging between pages 56-57 and pages 120-121.) A 
lesser annoyance is created by the absence of titles in some five percent of the 
bibliographic references. Finally, I note that at least one investigator whose work 
is critically discussed has been omitted from the author index. 

A few more substantive matters are of greater concern. The quality of some 
of the black-and-white photographs is quite poor (e.g. plates 10.4 and 14.6), and 
one wishes that better specimens could have been selected for certain illustrations 
(e.g. plates 10.5 and 13.2). With regard to the literature, a few recent papers 
have been overlooked, in particular those of Klots (1964-1968, J. N. Y. Entomol. 
Soc.) and others dealing with melanism in North American species. 

This book is a highly personal document, and as such must reflect the personality 
of its author. This reflection is generally engaging, and some occasional lapses of 
objectivity, particularly with regard to certain theoretical areas of biology, serve to 
enliven the sometimes tedious text. There are, however, certain dangers in this 
approach, and sometimes a question of fairness arises (e.g. with regard to 
ornithologists, p. 121). More serious, of course, would be any unfairness to specific 
individuals. On occasion, to this reviewer's mind, the data and conclusions of 
certain workers are disputed, re-interpreted, or rejected without an adequate airing. 

Elaboration of the following example may strike some as improper, but I risk 
that judgment in order to call attention to what I regard as a serious mis- 
representation. In the section of the book dealing with experiments on the 
background preferences of moths (p. 68-72), Kettlewell cites four papers of this 
reviewer, and details strong criticisms of the experimental techniques therein re- 
ported. He goes on to assert that in these studies “the main issue is missed,” i.e. 
whether the forms of polymorphic species differ in background preferences. In 
response, I must contend that the criticisms of techniques would only be ap- 
plicable had those techniques failed to yield readily interpretable results, and 
would point out that two polymorphic species (with a melanic form in each case) 
were tested for background preferences in the papers that are cited. (Kettlewell 
rather curiously overlooked another paper (1969, Nature, Lond. 222: 585-586) 
in which the background preferences of the typical and melanic forms of Phigalia 
titea were tested. ) 

This book is primarily, and properly, a vehicle for the elaboration of Kettlewell’s 
own ideas on all aspects of melanism, but one might have hoped for a fuller treat- 
ment in certain areas. For example, little treatment is accorded the possibility that 
various factors associated with industrialization, other than darkening of the en- 
vironment, might act, either directly on the insects, or indirectly through effects 
on predators or the vegetation, to provide an advantage to melanic individuals. 
Another idea which receives scant attention, and for which there is considerable 
experimental evidence, is the possibility that the melanic forms of cryptic species 


might differ genetically from their typical counterparts in terms of background 
resting preferences. 

In summary, while this book may fall somewhat short of expectation with regard 
to scholarship and synthesis, it is on the whole an ambitious and admirable project. 
Herein are compiled the results of two decades of substantial and varied investiga- 
tions by the author and his associates on the phenomenon of melanism in the 
British Isles. As a single source of these many results, this book will have a permanent 

THEODORE D. SarcENT, Department of Zoology, University of Massachusetts, 
Amherst, Massachusetts 01002. 

BUTTERFLIES OF THE WorLpD, by H. L. Lewis. 1973. Harrap Books, London; and 
Follett, Chicago. xvi + 312 p.; 208 pls. Price: about $30.00 (U:S.). 

No book could begin to live up to the pretentious title of this one, though in 
some respects Butterflies of the World makes a good attempt. There are recognizable 
figures of many (definitely not even most) of the world’s species, and the figures 
alone would make the volume worth far more than its purchase price if all of the 
species were correctly identified. 

Regrettably such is not the case. I have the feeling that Brig. Lewis prepared 
the text and the legends for the figures based on one idea of how the insects on 
each plate would be numbered, but that someone else did the final numbering. 
Those plates with even columns and rows of figures do not show transpositions, 
only those with irregularly placed specimens. Nevertheless, the presence of such 
easily avoided errors suggests careless proofreading and is inexcusable. <A partial 
listing of the plates affected by transpositions of numbers includes plates 19, 60, 63, 
64 and 118; there are otl.ers. Such errors greatly diminish the accuracy and useful- 
ness of the book. 

Errors of fact are even less excusable. Anartia amalthea (1L.), figured on Plate 
13 and listed from “N. and C. America,” is in fact a South American butterfly that 
has not been recorded from either North or Central America, though Seitz lists it 
without documentation from Central America. Troides aeacus (C. & R. Felder) from 
the Indo-Australian region is figured on Plate 24 as Eurytides xanticles (Bates) from 
the American tropics: even utilization of a rudimentary knowledge of Lepidoptera 
could have prevented this mistake. The genus Anetia has been variously considered 
a danaid (correctly) or simply a nymphalid, but not a heliconiid as figured on Plate 
43; and the danaid genus Ituna is included on Plate 44 as a heliconiid. Tellervo, 
the only Indo-Australian ithomiid, is shown on Plate 156 as a danaid. 

Lewis states (p. xii), “. . . the names given in the book are those commonly in 
use, and to be found in the latest works of scholarship. . . .” Unfortunately, the 
“latest work of scholarship” published in the Western Hemisphere seems to be 
Klots’ 1951 Field Guide! At the same time Forster’s Bolivian satyrid work is partially, 
but not critically, accepted, resulting in Altopedaliodes tena (Hewitson) being figured 
as that on Plate 54 and as Pedaliodes tena (Hewitson) on Plate 63. Some nomen- 
clatorial questions that were thought to be solved have been rescrambled, such 
as the distinction between Euphyes and Atrytone (Plate 21) and the replacement 
of the preoccupied Plestia by Zestusa (Plate 22). 

There is a small Corrigenda sheet accompanying our copy of this book: un- 
fortunately it should be much larger! The number of inadvertent synonyms created 
is very large (e.g., Mitoura spinetorum for M. spinetorum on Plate 20), and a 
full errata sheet should be forthcoming to rectify these errors. Since this book is 

VOLUME 28, NUMBER 2 179 

being merchandised by many booksellers throughout the English-speaking world it 
will be bought by many budding lepidopterists, ones who will never see a review that 
points up some of the errors in it. For the future accuracy of records provided by 
these people, an errata sheet (really a pamphlet) is not only desirable, but a must. 

This book is fairly good and accurate for the Old World and quite poor and 
out-of-date for the New. Perhaps one expects too much from a book that purports 
to be what this one does, but it simply is not a good book. I personally feel that 
Lewis was fighting a deadline and sacrificed final accuracy for a publication date. 
We would all be happier had he not! Nevertheless, if one takes the determinations 
with the proverbial “grain of salt,’ the figures make the book useful, for there are 
the best available representations of many poorly-known species contained within 
it. Further, one can get a general idea of what to expect in an area, even if he 
dare not rely on the names. 

Lee D. Mitter, Allyn Museum of Entomology, 3701 Bay Shore Road, Sarasota, 
Florida 33580. 

HEWITSON ON BUTTERFLIES, 1867-1877, with a Preface by Dr. L. G. Higgins. 1972. 
E. W. Classey Ltd., Hampton, Middlesex, England, iv + 242 p. (including cover 
pages). Price: $12.50 (U.S.). Distributed exclusively in the United States by 
Entomological Reprint Specialists, P. O. Box 77224, Dockweiler Station, Los Angeles, 
California 90007. 

This volume contains reprints of four privately published works by William 
Chapman Hewitson: “Descriptions of one hundred new species of Hesperidae” 
(1867-1868 ), “Descriptions of some new species of Lycaenidae” (1868), “Equatorial 
Lepidoptera collected by Mr. Buckley” (1869-1877) and “Bolivian butterflies col- 
lected by Mr. Buckley” (1874). In these remarkable papers Hewitson described 
ten new genera and no fewer than 403 new species of butterflies, all of which 
he intended to figure in either his Illustrations of . . . Exotic Butterflies (1852-1872) 
or Illustrations of Diurnal Lepidoptera (1863-1878). Regrettably Hewitson could 
figure only about half of the species described in these four pamphlet sets. 

L. G. Higgins’ Preface gives a brief, but interesting account of Hewitson’s lite 
and the histories of these important works; the works themselves give an insight 
into Hewitson. Hewitson was a wealthy enough man to afford the publication 
of his own papers and a proud one, as demonstrated by his comments justifying 
the publication of “Descriptions of some new species of Lycaenidae”: 

“Were I aware that any entomologist was engaged in a monograph of any 
particular group of butterflies, I should consider that I merely performed an act 
of common courtesy in avoiding said group until he had done with it. An 
entomologist, knowing that I am and have been for some time engaged in a 
monograph of the Lycaenidae, has, fortunately for me, given me notice that 
he is about to describe all those species in his possession. It is therefore in self- 
defence alone that I have been driven, greatly against my wish, to publish the 
following descriptions of species . . . 

Most systematists can identify with Hewitson in this complaint! 

While it was Hewitson’s fondest wish that he could figure all of the species 
described by him, he was unable to complete the task. Inasmuch as the types are 
preserved in the British Museum (Natural History), though some have not been 
identified with certainty, this volume could have been made an _ exceedingly 
valuable contribution by the inclusion of even black-and-white photographs of the 


relevant Hewitson types. Perhaps these photographs, along with a more modern 
treatment of the names, can be the basis of a future companion volume to the 
present one. 

The specialist can ill-afford to be without this book, unless he is fortunate enough 
to have access to the original papers. The amateur, however, can gain little from 
it because of the absence of illustrative material. The insights into Hewitson alone 
may make the book interesting to the general reader, especially if he is a history buff. 

Lee D. Mituer, Allyn Museum of Entomology, 3701 Bay Shore Road, Sarasota, 
Florida 33580. 

Published by the Amateur Entomologist’s Society, Feltham, Middlesex, England 
(A. E. S. Leaflet No. 34). 16 p., 15 figs. Price: not known. Available through 
the Publications Agent, L. Christie, 137 Gleneldon Road, Streatham, London, S. W. 
16, England. 

This little work is a fine introduction to the dissection and study of the genitalia 
of Lepidoptera, written clearly and concisely and well illustrated by line drawings 
of actual genitalia and the procedures for dissecting them. Simple step-by-step 
directions for the dissection of these structures make the study of genitalia some- 
thing for everyone, not a deep, mysterious subject restricted to the “experts.” 

The functions of the genitalic structures are discussed so that they have a biological, 
as well as taxonomic, significance for the amateur. The nomenclature of the parts 
follows Tuxen’s Taxonomists’ Glossary of the Genitalia in Insects (1956), the only 
text available which attempts homologies between genitalic structures in the various 

The mounting medium for the genitalia discussed is Euparal which is not in 
vogue on this side of the Atlantic, most slides being prepared here in Canadian 
balsam or synthetic equivalents. The only real difference in technique invoived is 
dehydration of the genitalia in 95% ethanol, then xylene, and final mounting in 
the balsam-type medium. 

The Amateur Entomologist’s Society is to be congratulated on preparing a very 
handy and informative little booklet. I wish I had an idea of the cost (I am certain 
it is nominal), but this information can be obtained from the Publications Agent, 
Mr. Christie. It would be worth doing so for anyone who wishes to become familiar 
with these important structures. 

Lee D. Miter, Allyn Museum of Entomology, 3701 Bay Shore Road, Sarasota, 
Florida 33580. 



Editor: THroporEe D. SARGENT, Department of Zoology, 
University of Massachusetts, Amherst, Massachusetts 01002 

K. S. Brown, J. M. Burns, R. H. Carcasson, J. P. DONAHUE, 
J. F. Gates Ciarke, C. D. Ferris, R. O. KENDALL, H. K. CLENCH, 
J. H. Masters, L. D. Miter, A. P. Puatt, A. M. SHApPio, J. R. G. 


Contributions to the Journal may deal with any aspect of the collection and study 
of Lepidoptera. Contributors should prepare manuscripts according to the following 

Text: Manuscripts should be submitted in duplicate, and must be typewritten, 
entirely double-spaced, employing wide margins, on one side only of white, 8% x 11 
inch paper. Titles should be explicit and descriptive of the article’s content, including 

the family name of the subject, but must be kept as short as possible. The first men- 

tion of a plant or animal in the text should include the full scientific name, with 
authors of zoological names. Insect measurements should be given in metric units; 
times should be given in terms of the 24-hour clock (e.g. 0930, not 9:30 AM). 
Underline only where italics are intended. References to footnotes should be num- 
bered consecutively, and the footnotes typed on a separate sheet. 

Literature Cited: References in the text of articles should be given as, Sheppard 
(1959) or (Sheppard 1959, 1961a, 1961b) and all must be listed alphabetically 
under the heading LirERaATuRE CirTeEp, in the following format: 

_ Sweprarp, P. M. 1959. Natural Selection and Heredity. 2nd. ed. Hutchinson, 

London. 209 p. 
196la. Some contributions to population genetics resulting from the 
study of the Lepidoptera. Adv. Genet. 10: 165-216. 
In the case of general notes, references should be given in the text as, Sheppard 

(1961, Adv. Genet. 10: 165-216) or: (Sheppard 1961, Sym. Roy. Entomol. Soc. 
London 1: 23-30). 

Illustrations: All photographs and drawings should be mounted on stiff, white 

: backing, arranged in the desired format, allowing (with particular regard to lettering) 
_ for reduction to their final width (usually 4% inches). Illustrations larger than 8% 
~ & 11 inches are not acceptable and should be reduced photographically to that size 

or smaller. The author's name, figure numbers as cited in the text, and an indication’ 
of the article’s title should be printed on the back of each mounted plate. Figures, 
both line drawings and halftones (photographs), should be numbered consecutively 

_ in Arabic numerals. The term “plate” should not be employed. Figure legends must 
_ be typewritten, double-spaced, on a separate sheet (not attached to the illustrations), 

headed EXPLANATION OF FicuREs, with a separate paragraph devoted to each page of 


Tables: Tables should be numbered consecutively in Arabic numerals. Headings for 

_ tables should not be capitalized. Tabular material should be kept to a minimum and 

must be typed on separate sheets, and placed following the main text, with the approx- 
imate desired position indicated in the text. Vertical rules should be avoided. 

Proofs: The edited manuscript and galley proofs will be mailed to the author for 
correction of printer's errors. Excessive author’s changes at this time will be charged 
to authors at the rate of 75¢ per line. A purchase order for reprints will accompany 
the proofs. 

Page Charges: Authors with grant or institutional funds are requested to pay a 

“a charge of $24.00 per printed page (including tabular and black-and-white illustrative 
_ material) for articles up to 20 pages in length. This charge may be waived in the 
- case of authors who have no grant or institutional funding, as it is not intended that 

any author should pay this charge from personal funds. However, all authors will 
be requested to pay this charge for material in excess of 20 printed pages. 
Address all correspondence relating to the Journal to the editor. Material not 

intended for permanent record, such as current events and notices, should be sent to 
_ the editor of the News: Ron Leuschner, 1900 John Street, Manhattan Beach, 
| California 90266. 




(Continued from outside front cover) 

EDITHA (NYMPHALDDAE). Raymond R. White and Michael C. 
Singer: ) ho ye a 

Nevapa. Thomas C. Emmel and John F. Emmel __ 


Scott L. Ellis 8 sn ee 

A New Species OF CoPTropiscA (HELIOZELIDAE) FROM MiIssissIpPi 
Jennings, Se ee 
Gapt: Michael J. Smith — 9. 
MeELANISM IN Motus or CENTRAL MassAcHuseTTs (NocTruméE, 
GEOMETRIDAE). Theodore D. Sargent __._______ 
(Nocrumae). D. H. Habeck, R. T. Arbogast and L. D. Cline 
DE CuBa, 1832. Roderick R. Irwin 0 ee 


A further field note on Isoparce cupressi (Sphingidae). Richard B. 
Dominick, is obs SE Ge, BR eg a ee 
Mortality in a group of Megathymus yuccae (Megathymidae). Ronald R. 
Gatrelle: °).52) 5 I PE 
Southern records of Mitoura hesseli (Lycaenidae). Richard A. Anderson 
An aberrant interspecific hybrid of Limenitis (Nymphalidae) from Wis- 
consin. Kuré Johnson, 220. 
Extended range distribution notes on Geometridae. Roger L. Heitzman __ 
A note on habitat and geography. Clifford D. Ferris __..______ 
Lepidoptera feeding at puddle-margins, dung and carrion. D. G. 
Sevastopulo (05 8 A ee 
Larval foodplants and parasites of some Lepidoptera in southeast Arkansas. 
Richard L.- Brown and Robert T. Allen _. "a 
A note on freeze-drying caterpillars. Richard S. Peigler = 
Freeze-drying and vacuum dehydration: instrumentation. Richard B. 
Dominick (2) eo aN 
The distribution and larval foodplant relationships of Saturnia walterorum 
(Saturniidae):’ P.M. Tuskes’<2200.° 0 

An unusually long pupal stage of Battus oldie polydamas L. mM 

(Papilionidae). Alberto Muyshondt |. 
An invasion of eastern Colorado by Vanessa cardui (Nymphalidae). F. — 
Martin “Browns 22080) eo 
Amphion nessus (Sphingidae) attracted to female Anisota virginiensis 
pellucida (Citheroniidae). Richard B. Dominick _.. 2 

Boox REVIEWS) 003.0 Cy Se ED a 

1974 Number 3 


of the 



: Appress—1973. Tur NatIOoNAL COLLECTION OF LEPI- 
er Capers erica onc ote Cue oN 181 

| ie 205 

erie re VU a 

(Continued on outside back cover) 

30 September 1974 


Harry K. CLencH (Pittsburgh, Penn.) President 

ANDRE BLANCHARD (Houston, Texas) President-elect 
RonaLtp W. Hopces (Washington, D.C.) Ist Vice President 
J. C. E. Riorre (Toronto, Ontario), Vice President 

L. Vari (Pretoria, South Africa) Vice President 

S. S. NicoLay (Virginia Beach, Va.) Treasurer 

Lee D. MILLER (Sarasota, Florida) Secretary 

Members at large (three year term): R. O. KENDALL (San Antonio, Tex.) 1975 
J. M. Burns (Cambridge, Mass.) 1974 J. A. PowEtu (Berkeley, Calif.) 1975 
R. H. Carcasson (Vancouver, B.C.) 1974 J. T. Brewer (Auburndale, Mass.) 1976 
M. C. NieEtsen (Lansing, Mich.) 1974 K. S. Brown (Rio de Janeiro, Brazil) 1976 
D. C. Fercuson (Washington, D.C.) 1975 K. W. Puiwie (Fairbanks, Alaska) 1976 

The object of the Lepidopterists’ Society, which was formed in May, 1947 and 
formally constituted in December, 1950, is “to promote the science of lepidopterology 

in all its branches, .. . . to issue a periodical and other publications on Lepidoptera, 

to facilitate the exchange of specimens and ideas by both the professional worker and 
the amateur in the field; to secure cooperation in all measures” directed towards 
these aims. 

Membership in the Society is open to all persons interested in the study of 
Lepidoptera. All members receive the Journal and the News of the Lepidopterists’ 
Society. Institutions may subscribe to the Journal but may not become members. 
Prospective members should send to the Treasurer full dues for the current year, 
together with their full name, address, and special lepidopterological interests. In 
alternate years a list of members of the Society is issued, with addresses and special 
interests. There are four numbers in each volume of the Journal, scheduled for 
February, May, August and November, and six numbers of the News each year. 

Active members—annual dues $10.00 
Student members—annual dues $7.50 
Sustaining members—annual dues $20.00 
Life members—single sum $150.00 
Institutional subscriptions—annual $15.00 

Send remittances, payable to The Lepidopterists’ Society, and address changes to: 
S. S. Nicolay, 1500 Wakefield Dr., Virginia Beach, Virginia 23455. 

Memoirs of the Lepidopterists’ Society, No. 1 (Feb. 1964) 

by Cyru. F. pos PAssos 

Price, postpaid: Society members—$5.00, others—$7.50; uncut, unbound signatures 
available for interleaving and private binding, same prices; hard cover bound, mem- 
bers—$8.00, others—$10.00. Revised lists of the Melitaeinae and Lycaenidae will be 
distributed to purchasers free (separately with paper covered copies and unbound 
signatures, bound in with hard covered copies). 

The Lepidopterists’ Society is a non-profit, scientific organization. The known 
office of publication is 1041 New Hampshire St., Lawrence, Kansas 66044. Second 
class postage paid at Lawrence, Kansas, U.S.A. 66044. 




Volume 28 1974 Number 3 


J. F. Gates CLARKE 

National Museum of Natural History, Smithsonian Institution, 
Washington, D.C. 20560 

James Smithson, an Englishman of noble birth, the natural offspring 
of the Duke of Northumberland and Elizabeth Keate Macie, a lineal 
descendant of Henry VII, bequeathed his estate to the United States 
Government “to found at Washington, under the name of the Smithsonian 
Institution, an establishment for the increase and diffusion of knowledge 
among men. Receipt of the bequest in 1838 precipitated a lengthy 
debate in Congress on whether the Government should, or indeed, could 
legally accept the funds and accompanying trust. In the Act of 1846, 
establishing the Smithsonian Institution, provision for a museum was 
made, and the name “United States National Museum” came into use 
in the year 1859. In 1884 appropriations to the Smithsonian for the U.S. 
National Museum were authorized and an annual report to the Congress 
by its Director was required. Today, the Museum’s component parts are 
the National Museum of Natural History and the National Museum 
of History and Technology. It is the former in which we are interested 

This brings us to the National Entomological Collections. The col- 
lections of all groups of insects consist of over 22,000,000 specimens. 
Of these approximately 3,500,000 are Lepidoptera. 

The National Collection of Insects is now over 93 years old having been 
started in 1881 with the transfer of 50,000 specimens to the U.S. Na- 
tional Museum from the U.S. Department of Agriculture. In 1886, C. V. 
Riley (1843-1895), then Chief Entomologist with the U.S. Department 



Fig. 1. C. V. Riley, 1843-1895 

VoLUME 28, NuMBER 3 183 

of Agriculture and honorary curator of entomology at the United States 
National Museum (appointed in 1882), donated 150,000 specimens of 
insects to the museum. In neither of the above cases was the number 
of Lepidoptera noted, indeed, it was not even certain that there were 
any Lepidoptera in these collections, but “C. V. Riley collection” labels 
are now occasionally found on specimens of Lepidoptera. Shortly before 
this time it is said that Townend Glover (1812-1883) also made a 
contribution to the National Museum, but there are no records in the 
Smithsonian to prove this claim. Since the earliest times the USS. 
Department of Agriculture’s entomological organizations and its entomol- 
ogists have been closely linked with the National Museum and, indeed, 
the Department of Agriculture’s contributions to the National Collection 
have been great and many. 

There was no paid entomologist on the Smithsonian’s staff until 1885 
when John B. Smith, a lepidopterist, was appointed curator. From 1900 
to 1940 the Smithsonian had one staff entomologist. This has now 
increased to 12, four of whom are lepidopterists. The U.S. Department 
of Agriculture had 16 specialists in 1930 and 29 in 1974. Of these, 
four are lepidopterists. 

Five men, William Schaus, August Busck, Carl Heinrich, H. G. Dyar 
and William Barnes (the latter not a member of the museum staff) were 
primarily responsible for the early development of the National Col- 
lection of Lepidoptera. 

August Busck (1870-1944) 

August Busck was born in Randers, Denmark, 18 February 1870 and 
in 1893 he came to the Columbian Exposition in Chicago. In March 
1896 he was appointed assistant in the Division of Entomology, U.S.D.A., 
becoming shortly thereafter a specialist in Microlepidoptera. He investi- 
gated the mosquito and Lepidoptera faunas of the West Indies in 1905 
under the auspices of the Carnegie Institution and made a similar 
investigation of the mosquito and Lepidoptera faunas of the Canal 
Zone for the Panama Canal Commission in 1907. Parts of the results 
of these two expeditions were reported in the Howard, Dyar, Knab 
monograph of The Mosquitoes of North and Central America and the 
West Indies. Most of the Lepidoptera collected by Busck were reported 
by Dyar and himself, but some of Busck’s material is still being used 
by today’s workers. 

Busck was one of the first to use consistently genitalic characters, 
the most important structures ever to be used in the classification of 
the various categories of Microlepidoptera. 


1) Fig. 2. John B. Smith, 1858-1912 


VoLuME 28, NUMBER 3 






Fig. 3. August Busck, 1870-1944 


Throughout his career Busck was intimately associated with the 
USNM and its collection of Microlepidoptera is the second most ex- 
tensive and important in the world, and for this eminence Busck must 
be given chief credit. 

Harrison Gray Dyar (1866-1929) 

Harrison Dyar was born in New York City and was educated at 
the Massachusetts Institute of Technology, Columbia University and in 
the field. He came to the U.S. National Museum in 1897 but was on 
the payroll of the U.S.D.A. 

He was an extremely versatile man. He had a good eye for species, 
could comprehend major groups of insects and had the ability to study 
intensively the biology and early stages of one or another group. He 
excelled in all these fields and was able to make a synthesis of them. 
His publications include a larger number of full and accurate larval 
descriptions than the work of any other American entomologist. On 
the basis of his knowledge of adults, larvae and eggs his work may per- 
haps be considered the basis of our modern classification of moths. 

His greatest work, however, in which he did the taxonomy, was the 
monumental work The Mosquitoes of North and Central America and 
the West Indies published in four volumes (1912-1917) by the Carnegie 
Institution. He is also the discoverer and promulgator of what has 
become known as “Dyar’s Law’—that the widths of the head of a 
larva in its successive stages follow a regular geometrical progression. 

There was another entomological law that emanated from the National 
Museum known as “Schwarz’s law.” This refers to the accidental oc- 
currence of an insect on a plant and thus the misinterpretation of its 
host and reminds us that “a bug must have some place to sit.” 

Wilhelm Carl Paul Gottlieb Heinrich (1880-1955) 

Carl Heinrich was born in Newark, New York, 7 April 1880 
and has been described as a poet, writer, student of music, history 
literature and philosophy. He intended to study music under Edward 
McDowell but before the opportunity came McDowell died. He came 
to Washington in 1902 but did not begin working with the U.S. De- 
partment of Agriculture until 1913 and retired his services in 1949. 

He published 87 papers and books, many of them editorial in nature. 
His two largest works were the Revision of the family Olethreutidae 
in two parts (1923-1926) and American moths of the subfamily Phycitinae 
(1956). His publications were not confined to entomological subjects. 

< ¢ « 


pleAroTZ WIeIT[IAA “VAC “s) UOSELIBET :7YStI 0} 4yZo"T ‘Sly 

















Fig. 5. Carl Heinrich, 


eS ee Se et ae a a a 

VOLUME 28, NUMBER 3 189 

In 1901 he published a book of poems Moods and Moments and in 
1929 his controversial, satirical The Orphan of Eternity or the Katabasis 
of Lord Lucifer Satan. In retirement he wrote 105 articles on subjects 
of public interest published in the News and Courier of Charleston, 
South Carolina. 

William Schaus (1858-1942 ) 

William Schaus was born in New York City 11 January 1858, the 
son of a well known art collector and dealer. He was schooled largely 
in Europe and his principal training was in music, art and languages; 
but as a young man he came under the influence of Henry Edwards and 
found his real vocation, in spite of paternal opposition. 

Schaus made his first important collecting trip to Mexico in 1883 and 
subsequently made frequent and extended trips, with his companion 
Jack Barnes, to Mexico, Costa Rica, Guatemala, Panama, Cuba, Jamaica, 
Dominica, St. Kitts, the Guianas, Colombia and Brazil and collected 
over 200,000 Lepidoptera. He was one of the great contributors to, 
and dedicated workers in, the Lepidoptera collection. Schaus sums up 
his attitude to the National Collection in a letter from Costa Rica dated 
15 March 1909, to Richard Rathbun, then Assistant Secretary of the 
Smithsonian, in charge of the National Museum, “I . . . announce to you 
the gift of my butterflies and Sphingidae to the Museum, as they were 
not included in the large collection of moths I gave the Museum three 
years ago—lI am glad to be able to do so—I am still hard at work and 
securing many new and rare species, so there is no danger of the 
Smithsonian losing its foremost place as possessor of the finest collection 
of Tropical American Lepidoptera.” 

Schaus came to the National Museum in 1895. By 1906 he was back 
in Mexico. An accession of 7 June 1906 records “Large collection of 
Lepidoptera.” Schaus’ letter accompanying the gift states “The box 
contained . . . 22 parcels of Lepidoptera.” Other accessions merely 
list “1 box Lepidoptera in papers (1907) Costa Rica, or “291-cartons 
of unmounted insects, Costa Rica.” Other accessions record material 
from Argentina, Guatemala, Bolivia, Ceylon, New Guinea, Ecuador and 

From 1919 until his retirement in 1938, Schaus was on the staff of 
the U.S. Department of Agriculture and in 1921 was made honorary 
assistant curator of insects of the U.S. National Museum. During his 
lifetime he published 122 papers in which he described more than 
5,000. species. 


if Fig. 6. William Schaus, 1858-1942 

VOLUME 28, NUMBER 3 191 

William Barnes (1860-1930) 

William Barnes was born in Decatur, Illinois, and except for his 
medical training at Harvard, post graduate medical study in Germany 
and some travelling, spent all his life there. He brought together the 
greatest, finest and most complete and most accurately determined 
collection of North American Lepidoptera in the world. If Dr. Barnes 
had done nothing but assemble this collection he would have done 
a great work, but he did much more. His copiously illustrated Con- 
tributions to the natural history of the North American Lepidoptera, 
which occurs in 5 volumes, embodies the researches of himself and his 
collaborators, and consists of extensive descriptive and revisionary papers. 

Some details about the Barnes collection follow: 

As early as 1921 a move was afoot to obtain for the National Museum 
the William Barnes collection. J. M. Aldrich, Associate Curator of 
insects, stated in a letter of 31 October 1921 to Dr. Stejneger, Head 
Curator of Zoology: “The value of the collection is so great that we 
can make no offer from the current revenues of the National Museum 
or the Smithsonian, that would not be absurd and pitiful. The case will 
require special action by Congress, I should think.” Dr. Barnes was 
seeking $200,000. What he had actually proposed doing was to give 
the collection to the Decatur and Macon County Hospital to be sold by 
them for added revenue. In 1922, Representative Moore of Illinois 
presented a bill (H.R. 10597) asking for enactment of authorization for 
the Secretary of the Smithsonian to purchase the Barnes collection for 
$310,000 (including shipping). Negotiations continued until Dr. Barnes’ 
death in 1930. The late Secretary Abbott, of the Smithsonian, on June 
13, 1930 sent a memo to Dr. Wetmore, then Director, U.S. National 
Museum, stating “The executors desired to give the National Museum 
first opportunity to obtain the collection for $50,000. (Barnes is reputed 
to have spent $400,000 on the collection.) The collection was finally 
purchased through a congressional appropriation with a rider on an 
agriculture deficiency bill. Carl Heinrich and August Busck prepared 
the collection for shipment and the collection was accessioned February 
iG 1931. 

By the time I arrived at the National Museum both Barnes (who 
was not on the staff) and Dyar were dead, but Busck, Heinrich and 
Schaus were still active. 

For many years Busck and Heinrich performed like twins. They 
dressed alike, both used a cane, and about the only way they could 


Fig. 7. William Barnes, 1860-1930 

VOLUME 28, NUMBER 3 193 

be distinguished at a glance was by the fact that Heinrich wore a monocle. 
This close relationship went on for years but finally something happened, 
to which I was not privy, and the friendship ended. After that, even 
up to the time of Busck’s retirement in 1940 this feud lasted. They were 
civil enough to each other at the office, but after official hours there 
was no communication between them at all. When Busck’s end was 
near, early in 1944, Heinrich relented and decided to go and visit, for 
the last time, this old friend. Busck was on his death bed, but said 
to Heinrich, “Vell, Carl, let's go downstairs and have a drink, I don't 
vant to die sober!” Busck died shortly thereafter. 

For many years it was customary to accumulate reprints in the 
Division of Mammals, where we were allowed to examine and select 
what we wanted for our files. The practice was to indicate a_ bid, 
usually 5¢, and initial the amount offered. If someone wanted the 
reprint more than the first bidder he increased the offer to 10¢, and 
so on until bidding was closed. During one of these “auctions” a copy 
of Busck’s A Revision of the American Moths of the Family Gelechiidae . .. 
1903, appeared. I did not have this paper so I offered the usual 5¢. 
Busck’s pride was hurt; he was angry. He burst into my office and 
roared, “I have just seen your bid on my gelechiid paper. I have bid 
25¢ and if you vant the paper you will have to pay 30¢!”. I did. 

Heinrich is reputed to have started each day with a martini! He smoked 
cigars and cigarettes incessantly and used snuff. The smokers in the 
museum were provided with spittoons for their cigar and cigarette ashes 
and butts up until the 1950's. The spittoons were also used to throw 
waste from dissections. One day, inadvertently, when Heinrich threw 
away some waste from a dissection an aedeagus from a type went with 
it. Obviously, that was a loss that could not be accepted so Heinrich, 
with an aide, spent the day going through cigar butts, cigarette tobacco 
and waste, spoonful by spoonful, looking for the aedeagus. They found 

Schaus was a bachelor. On one of his visits to England he employed 
a valet, Jack Barnes, who became his lifelong companion. Early in their 
association Schaus, in admiration for his friend, bought him a quantity 
of stocks. In the crash of 1929 Schaus lost much but after the crash 
Barnes’ stocks appreciated and the valet became wealthier than the 
master. When Schaus died he left Barnes a considerable amount of 
money, plus a valuable stamp collection so Barnes was fixed for life. 

Schaus enjoyed a wide correspondence throughout much of the world 
and wrote his correspondents in English, German, French and Spanish, 
as appropriate. 


Us Species. 

Fig. 8. A drawer containing Zephy? 

VOLUME 28, NUMBER 3 195 

It is no wonder that Schaus described 5000 species. He took his work 
home with him! Nearly every night he would take home with him a 
small box containing half-a-dozen specimens, each representing a dif- 
ferent species. In the morning he would return with each species de- 
scribed and each specimen labeled “type.” Over the years there were 
some slight deviations from this practice but one can generally depend 
on the specimens labeled “type” as the ones he carried home. 

Schaus had many friends among the wealthy and it was through 
them that he, singlehandedly, raised the money for the purchase of the 
Dognin collection. Schaus told me that the purchase price was over- 
subscribed and that he had had to return checks of $5,000 and $10,000. 

In preparation for his retirement Schaus was cleaning out all personal 
belongings, among them many reprints of his papers. I discovered that 
he was discarding them and decided to retrieve them for future distribu- 
tion. Schaus discovered this. One morning, not even stopping to remove 
his coat and hat, he stormed into my office and accused me of saving 
the reprints for later sale. From then on he tore each reprint in two 
before throwing it into the wastebasket! 

The museum collection of Lepidoptera is housed in approximately 
30,000 drawers, nearly 3,000 of which are devoted to Microlepidoptera. 
In recent years the unit tray system has been employed although the 
whole collection has not yet been converted to that type of housing. 
For large species, most Sphingidae, Saturniidae, Papilionidae, etc., whole 
drawers are still used, with no division being made in the drawers by 

Some of the more important accessions that are included in the na- 
tional collection are as follows: 

Alfieri, Anastase. 1966. (purchase). This material consists of Egyptian 

Heterocera, including many types. 

Baker, Charles Fuller. 1928 (gift). No breakdown of the more than 
300,000 specimens in this collection is given but it contains perhaps 
the largest number of Philippine Lepidoptera yet assembled by one 
person plus bountiful material from Malaya. 

Barnes, William. 1931 (purchase). The accession shows a record of 
473,500 but the actual count was 473,293 specimens including 1950 
holotypes. This collection consists mainly of North American ma- 
terial. In the Barnes collection is incorporated material from the 
Oberthtir, Taylor, Kearfott, Polling, Lacy, Field, Hill, Longley, 
Spalding and Merrick collections. 



Saawta Menkes exRTE IS 


mecdens HRTTIEY SRL 

VoLUME 28, NUMBER 3 197 

Fig. 10. Douglas Ferguson (USDA) examining specimens in one of the lanes 
of the range. © 


Blackmore, E. H. 1937-39 (gift). Though small, this collection is 
all from British Columbia and consists of 6950 specimens. 

Box, Harold. 1963 (gift). This collection consists of 5000 specimens, 
entirely of the important sugar cane feeding genus Diatraea. 

The Brighton Museum, Brighton, England. 1949 (gift). There are 
15,000 Microlepidoptera in this material. Nearly all species from 
England are represented. 

The Brooklyn Museum. 1929 (gift). This material consists of 37,000 
miscellaneous insects, and although the collection is comprised 
largely of Lepidoptera, the exact number of specimens of this order 
was never recorded. Contains types of Neumogen, Hulst and others. 

Clarke, J. F. Gates. 1937 (gift). From the Pacific Northwest. The 
original gift consisted of over 10,000 specimens. All Clarke types 
(over 300), except one, are in the national collection. 

Dognin, Paul. 1925 (purchase). This collection consists of 82,000 
specimens among which are 3,000 Dognin types and over 300 Thierry- 
Mieg types. The collection was purchased through public subscription 
and was then presented to the museum. 

Dyar, H. G. 1903 et seq. (gift). The first contribution recorded from 
Dyar consisted of 20,320 specimens from British Columbia. In 1917 
he added another 17,000 North American specimens and _ subse- 
quently numerous smaller gifts. 

Engelhardt, George P., 1941-1943 (gift). Contains over 9,000 
Aegeriidae. Its great value lies in the fact that nearly all specimens 
are reared and that larvae are associated with the adults. 

Ferguson, Douglas. 1970 et seq. (gift). There are 48,000 specimens 
from the Northeastern United States and eastern Canada, pre- 
dominantly from Nova Scotia; also from Newfoundland. 

Fernald, C. H. 1924-25 (purchase). This is a type collection of 
Microlepidoptera containing not only Fernald’s tortricid types, but 
types of Fitch’s Pterophoridae and types from Fish. The collection 
also contains cotypes of Walsingham, Hulst, Packard and Grote. 

Field, W. D. 1947 (gift). This gift of 5,000 specimens is composed 
of Japanese and European Rhopalocera. Field contributed numerous 
other smaller gifts of North American Lepidoptera. 

Graham, David C. 1918-1948 (gift). Over the many years indicated, 
the Rev. David C. Graham sent thousands of Chinese Lepidoptera 
to the National Museum. These were never counted, but were 
recorded as so many packages. 



Fig. 11. Donald Davis on Microlepidoptera range. 



VoLUME 28, NUMBER 3 201 

Hayward, H. C. 1949 (gift). The Hayward collection of English 
Tortricidae was obtained by the British Museum (Natural History ) 
and presented to the USNM. 

Hodges, Ronald W. 1962 (gift). Consists of 25,000 specimens 
primarily of North American Microlepidoptera largely from Arizona, 
New York and Florida. 

Hopfinger, J. C. 1962 (purchase). This collection is worldwide in 
scope but the preponderance is from Washington State. 

Jones, Frank Morton. 1950-56 (gift). Of the 8,460 insects given to 
the National Collection the most important segment consisted of 
4.400 specimens of Psychidae (bag worms). Most of the species 
included are from the Americas. 

Issiki, Syuti. 1972 (purchase). The Issiki collection constitutes the 
most complete assemblage of Japanese and Formosan Microlepidop- 
tera ever brought together and contains about 95 per cent of the 
known Japanese species. There are 78 holotypes and more than 
200 secondary types in this material. 

Meadows, Don. 1950 (purchase). There are nearly 9,000 specimens 
primarily from the Channel Islands, off the coast of California, in 
this accession. 

McAlpine, W. S. 1972 (gift). This collection consists of more than 
12,000 specimens, predominantly of the butterfly genus Calephelis, 
but also it is strong in miscellaneous Michigan Lepidoptera. Most 
of McAlpine’s types are included. 

McElvare, Rowland R. 1967 (gift). In this accession there are over 
4,200 specimens of the sub-family Heliothinae (Noctuidae) from 
North America. Types of McElvare’s species are in the National 

Nawa, U. 1903 (gift). Japanese Lepidoptera exhibited at the St. Louis 
World Fair constitute this gift. 

Philpott, A. 1928 (gift). Donated in this gift is a nearly complete 
collection of New Zealand Microlepidoptera. 

Rawson, George W. 1962-1972 (gift). The more than 9,000 specimens 
are mostly North American, predominantly from Michigan and 

Schaus, Wm. 1901 et seq. (gift). Altogether, as correctly as the records 
can be interpreted, Schaus donated most of approximately 200,000 
specimens he collected from the Neotropical Region. But many of 
his contributions were merely recorded as “3 crates” or “2 crates,” 
and the numerous specimens were never counted. About 5,000 
of his types are included. 



oe — 

1343 Bpai 


VoLUME 28, NuMBER 3 203 

Schonborn, Wm. E. and Theresa F. 1925 (gift). This collection con- 
sists of material from the eastern United States and Europe. 

Shoemaker, Ernest. 1957 (gift). This accession records 60,338 speci- 
mens of insects but is not broken down by order. The collection 
contains mostly Neotropical and Nearctic species and is rich in 
the genus Morpho. 

Smyth, J. Adger. 1947 (gift). Mr. J. Adger Smyth’s father, Dr. Ellison 
A. Smyth, made this collection of more than 16,000 Lepidoptera, 
including two types. It is worldwide in coverage. Subsequently 
(1970) J. Adger Smyth donated an additional 1,174 specimens from 
the Americas and Africa. 

Vallins, F. T. 1971 (purchase). This entire collection of 22,000 speci- 
mens of Lycaenidae is of Palaearctic origin. The collector en- 
deavored to assemble series of species from throughout their ranges, 
thus showing ail variations known. 

In addition to the collections obtained from gifts, purchases, and 
transfers from other governmental agencies, much accrues to the national 
collection through field work by a very active staff. In recent years 
there have been many expeditions to many parts of the world which 
have produced Lepidoptera. Extensive field work has been conducted 
in many parts of North America, nearly every country in Central and 
South America, and the West Indies, Australia, Africa and Borneo. On 
the island of Dominica (BWI) alone a survey of the terrestrial arthropods 
continued over a period of three years, with a change of team and 
emphasis every three months. A continuing program, now in its fifth 
year, is being conducted in Sri Lanka (formerly Ceylon). Much ma- 
terial has been acquired from the islands of the Western and South 
Pacific, including Micronesia, Society Islands, Tubuais, Marquesas and 
the Philippines. 

Copies and original photographs for this article have been made by 
Mr. Victor Krantz, staff photographer, Smithsonian Institution. 


AupricH, J. M. 1929. Entomol. News 40(5): 167-168. 

Dyar, H. G. 1890. The number of molts of lepidopterous larvae. Psyche 5(175- 
176): 420-422. 

Forses, Wm. T. M. 1929. Harrison Gray Dyar. Entomol. News 40(5): 165-167. 

Gunper, J. D. 1929. North American Institutions Featuring Lepidoptera. VIII. 
U.S. National Museum, Washington, D.C. Entomol. News 40(9): 281-286. 

HermnricoH, C. & E. A. CHapriin. 1942. William Schaus. Proc. Entomol. Soc. 
Washington 44(9): 189-195, pl. 17. 

. & U. C. Lorrin. 1944. August Busck. Proc. Entomo:. Soc. Washington 

AG(9): 231-239. 


Taytor, F. A. 1966. Museum of Natural History, U.S. National Museum. 1966 
Annual Report. 37-151, I-LVIII. 

True, W. P. 1946. The first hundred years of the Smithsonian Institution. I-VI, 
1-64, illus. Washington, D.C. 

VAN Duzer, E. P. 1930. Dr. William Barnes. Pan Pacific Entomol. 7(1): 16. 

Wane, J. S. & H. W. Capps. 1955. Carl Heinrich. Proc. Entomol. Soc. Washington 
BS) eo BE 55. 

WituraMs, R. C. & P. P. Catverr. 1942. Dr. William Schaus. Entomol. News 53: 


Pupae of Eupithecia die even when reared under what seem to be optimal condi- 
tions. In many cases considerable time is spent on descriptions of larvae with the 
expectation of rearing adults to enable one to determine the species. When the 
specimen dies in the pupal stage this time may be lost unless one can determine 
the species from the pupa. Sometimes pupal development is such that adult 
features can be seen within the pupal case. 

Method. To determine if the genitalia have developed sufficiently, sever the 
pupa between the fourth and fifth abdominal segments. If the internal organs have 
not developed sufficiently the abdomen will appear empty and it is of little use 
to proceed further. If such is the case, place the two halves of the pupal case 
in a gelatin capsule of suitable size and replace in the collection for future study. 
A shrivelled abdomen may, however, still be satisfactory for further work. About 
three-quarters of the specimens examined have had the genitalia developed suf- 
ficiently for one to make specific determinations. If the specimen seems developed 
enough, the entire severed abdomen is immersed in a 10 percent solution of potassium 
hydroxide for approximately 16-20 hrs. Do not attempt to forcibly remove the 
abdomen from the pupal case before imersion in KOH solution, unless it is already 
loose, otherwise both may be damaged. After removal from the caustic, place in a 
solution of 30 percent alcohol and the abdomen will separate very easily from the 
pupal case without damage. The empty portion of the pupal case should be placed 
out to dry and later put in a gelatin capsule along with the remainder of the pupal 
case. The cremaster and other important diagnostic characters can still be used for 
study. Process the abdomen as suggested by Hardwick (1950, Can. Entomol. 83: 
231-235). Extreme care should be taken, however, during the dissection as the 
material is usually much more fragile than in a fully mature adult. One should also 
keep in mind that the pupal case may contain a parasite which has died before 
emerging. These are often quite large and can be mistaken for a moth before 

This technique has proved useful for the determination of species in several 
other genera of Geometridae besides Eupithecia. Among these are the genera 
Deilinia Hbn., Rheumaptera Hbn., Hydriomena Hbn., Drepanulatrix Gump., 
Semiothisa Hbn., and Itame Hbn. This technique could very well prove useful in 
other families of Lepidoptera as well. 

K. B. Botre, Seconded from the Canadian Forestry Service, Department of the 
Environment, Ottawa, Ontario. 

VoLUME 28, NUMBER 3 205 


Department of Biology, Carleton University Ottawa, Ontario, Canada K1S 5B6 

Department of Biology, Tufts University, Medford, Massachusetts 02155 

Acoustic communication between moths and bats has provided biolo- 
gists with several intriguing examples of predator-prey interactions. 
Moths of some families, notably the Arctiidae, Ctenuchidae, Geometridae, 
and Noctuidae, possess tympanic organs capable of detecting the 
echolocating cries of bats (Eggers, 1928; Schaller & Timm, 1950; Haskell 
& Belton, 1956; Roeder & Treat, 1957). When presented with a source 
of ultrasonic pulses (cries of an echolocating bat or facsimiles thereof), 
restrained noctuid moths show changes in wingbeat and the same moths 
while flying free in the field perform evasive maneuvers (Treat, 1955; 
Roeder, 1962, 1967 a, b). 

When confronted with ultrasonic pulses, some arctiid and ctenuchid 
moths produce ultrasonic sounds usually consisting of a repeating 
sequence of brief, high-pitched, rapid clicks. These clicks are produced 
by a row of minute ridges or corrugations (= microtymbals or the 
‘striate band’ of Forbes & Franclemont [1957]) in the cuticle of the 
metathoracic episternite (Fig. 1). The basalar muscle inserts on the 
concave inner surface of the episternite, and its contraction cycle causes 
the microtymbals to buckle in sequence; each ‘buckle’ producing a click 
considerably less than one millisecond in duration and containing ultra- 
sonic frequencies (Blest et al., 1963; Blest, 1964). The contraction cycle 
of the basalar muscle determines the duration of the click cycle, and 
the number and form of the microtymbals on the episternite affects 
the number and acoustic nature of the clicks in each cycle. 

Some arctiids and ctenuchids are not only noisy, but often distasteful 
(Beebe & Kenedy, 1957; Rothschild, 1961; Blest, 1964; Rothschild & 
Alpin, 1971). Some arctiids evert cervical glands when threatened (eg. 
Utethesia ornatrix bella (L.); Eisner, 1970, Fig. 6a), and this may be 
accompanied by noise (eg. Arctia caja (L.); Rothschild, 1965). Potential 
predators of these species may learn to associate the noise with an 
unpalatable taste and thus use the noise as a signal to leave the moths 

Captive Myotis lucifugus (LeConte) (Chiroptera: Vespertilionidae) 
avoided noisy Haploa clymene (Brown), Halysidota tessellaris (Abbot 


Fig. 1. Diagram of left lateral wall of meso and metathorax of a noctuid moth 
with the metathoracic episternite heavily outlined in black and the wing area 
indicated by lines (epm = epimaron; eps = episternite; sa = subalar plate; scu = 
scutum ). 

and Smith), and Pyrrharctia isabella (J. E. Smith), but bit into 
them if they were quiet. However, having tasted the moth, the bats 
spat out the H. tessellaris and H. clymene, and ate the P. isabella 
(Dunning, 1968), apparently a Batesian mimic of the other two species. 
Coutts et al. (1973) observed that captive M. lucifugus and Eptesicus 
fuscus (Palisot de Beauvois) (Chiroptera: Vespertilionidae) bit into and 
then rejected dead or quiet Cycnia tenera Hubner. 


Fig. 2. Scanning electron micrographs of the left (except d and g) metathoracic 
episternites of: (a) Euthisanotia unio Hubner (Concord, Massachusetts); (b) 
Phragmatobia assimilans Newman and Donahue (Edmonton, Alberta); (c) 
Phragmatobia fuliginosa L. (Suisse Valias); (d) Hypoprepia fucosa Hubner (Con- 
cord, Massachusetts); (e) Hypoprepia cadaverosa Strecker (Greer Rd, White 
Mountains, Arizona); (£) Hypoprepia miniata (Kirby) (Attons Lake, Saskatchewan); 
(g) Clemensia albata Packard (Bobcaygeon, Ontario); (h) Turuptiana permaculata 
Walker (Buena Vista, Colorado); (i) Spilosoma congrum (Walker) (Lac Mondor, 
Quebec). Scale indicates 1 mm. 


VoLUME 28, NuMBER 3 

5 Ss : 
fA ES: 


Dunning & Roeder (1965) showed that flying M. lucifugus veered 
away from their prey (moths or ‘tossed mealworms’) when confronted 
with tape-recorded microtymbal sounds as they closed with their prey. 
However, one of Dunning’s (1968) captive bats learned to distinguish 
between the noises of P. isabella and H. clymene and H. tessellaris indi- 
cating that different moths produce different noises. 

These behavioural considerations and the possibility of marked species 
differences in microtymbal systems of arctiids led us to make a pre- 
liminary survey to determine the incidence of microtymbals in this group. 
The following is a brief presentation of some of our anatomical findings 
which may be of interest to lepidopterists. 

Figs. 2 and 3 are scanning electron micrographs of the metathoracic 
episternites of specimens obtained from collections at the Entomology 
Research Institute, Canada Department of Agriculture, and Carleton 
University. It is evident from the micrographs that the details of the 
microtymbals vary considerably both within and between genera. Sexual 
dimorphism, if it occurs, was not investigated. The metathoracic 
episternite of a noctuid, Euthisanotia unio Hubner, is shown for com- 
parison (Fig. 2a). 

Well developed microtymbals occur on the metathoracic episternites 
of the following species: Phragmatobia assimilans Newman and 
Donahue (Fig. 2b), Phragmatobia fuliginosa L. (Fig. 2c), Phrag- 
matobia lineata Newman and Donahue, Hypoprepia fucosa Hubner 
(Fig. 2d), Hypoprepia cadaverosa Strecker (Fig. 2e), Hypoprepia 
miniata (Kirby) (Fig. 2f), Clemensia albata Packard (Fig. 2g), Cisthene 
nexa Boisduval, Crambidia casta Sanborn, Crambidia pallida Packard, — 
Halysidota tessellaris (Fig. 3a), Halysidota maculata (Harris), Hemi- 
hyalea labecula Grote (Fig. 3d), Pyrrharctia isabella (Fig. 3f), Cycnia 
tenera (Fig. 3g), Euchaetias egle (Drury) (Fig. 3h), and Utethesia 
ornatrix bella (Fig. 3i). 

The following species have poorly developed microtymbals: 
Turuptiana permaculata Walker (Fig. 2h), Estigmene acrea (Drury), 


Fig. 3. Scanning electron micrographs of the left (except c, e, and h) metathoracic 
episternites of: (a) Halysidota tessellaris Abbot and Smith (Chaffey’s Locks, 
Ontario); (b) Halysidota argentata Packard (Nanaimo, British Columbia); (c) 
Halysidota caryae (Harris) (Normandale, Ontario); (d) Hemihyalea labecula Grote 
(Durango, Colorado); (e) Haploa clymene (Brown) (Arrowhead Lake, Myrtle Beach, 
South Carolina); (f£) Pyrrharctia isabella (J. E. Smith) (Lac Mondor, Quebec); 
(g) Cycnia tenera Hubner (Chaffey’s Locks, Ontario); (h) Euchaetias egle (Drury) 
(Concord, Massachusetts); (i) Utethesia ornatrix bella (1.) (Punta Gorda, Florida). 
Scale indicates 1 mm. 




Spilosoma congrum (Walker) (Fig. 2i), Arachnis maia Ottolengui, 
Spilosoma virginicum (Fabricius), Arctia caja, Platarctia parthenos 
(Harris), and Halysidota caryae (Harris) (Fig. 3c), and in Holomelina 
ferruginosa (Walker) and Halysidota argentata Packard (Fig. 3b), they 
are absent. 

The details of the surface of the microtymbals of U. o. bella (Fig. 31) 
are strikingly different from those of the other arctiids mentioned. 

Forbes & Franclemont (1957) considered using the ‘striated band’ of 
the Arctiidae as a taxonomic character but noted that it varied markedly 
in species considered on other grounds to be closely related. Their 
finding is strikingly confirmed by an examination of Figs. 2 and 3. 
Thus, the two species of Phragmatobia (Fig. 2b and c) and the three 
species of Halysidota (Fig. 3a, b, and c) show marked differences in 
the form and degree of development of the microtymbals. At the same 
time, the three species of Hypoprepia (Figs. 2d, e, and f) have micro- 
tymbals which are quite similar in form and arrangement. It will be 
interesting to see if microtymbal morphology correlates closely with other 
characteristics used in the classification of arctiids. 


We thank Drs. E. G. Munroe and E. A. Arnason for providing us with 
the specimens. We are especially grateful to Mr. L. E. C. Ling who took 
the micrographs of the uncoated specimens at low voltage using the 
techniques described by Howden & Ling (1973). We are also grateful 
to Drs. E. G. Munroe and H. F. Howden for critically examining the 
manuscript and to Dr. E. G. Munroe and Rev. J. C. E. Riotte for en- 
couraging us in this survey. Rev. Riotte kindly checked and corrected 
the names of the moths. This study was supported by National 
Research Council of Canada Operating and Equipment Grants to MBF 
and by a Career Award from the National Institute of Health, United 
States of America, to KDR. 


BEEBE, W. & R. Kenepy. 1957. Habits, palatability and mimicry in thirteen 
ctenuchid moth species from Trinidad, B.W.I. Zoologica 42: 147-157. 

Buiest, A. D. 1964. Protective display and sound production in some new world 
arctiid and ctenuchid moths. Zoologica 49: 161-181. 

, T. S. Cotter & J. D. Pyr. 1963. The generation of ultrasonic signals by 
a New World arctiid moth. Proc. Roy. Soc., London (B) 158: 196-207. 

Courts, R. A., M. B. FENton & E. GLEN, 1973. Food intake by captive Myotis 
lucifugus and Eptesicus fuscus (Chiroptera: Vespertilionidae). J. Mammal. 
54: 985-990. 

Dunninc, D. C. 1968. Warning sounds of moths. Z. Tierpsychol. 25: 129-138. 

VOLUME 28, NUMBER 3 211 

. & K. D. Roeper. 1965. Moth sounds and the insect-catching behavior of 
bats. Science 147: 173-174. 

Eccers, F. 1928. Die Stiftfuhrenden Sinnesorgane, Morphologie und Physiologie der 
chordotonalen und der tympanalen Sinnesapparate der Insekten. Zool. Bausteine 
2(1), 354 p., Berlin. 

Eisner, T. H. 1970. Chemical defense against predation in arthropods, in Chemical 
Ecology, E. Sondheimer & J. B. Simeone, eds., Academic, New York and 
London, p. 157-217. 

Forses, W. T. M. & J. G. FrANcLEMONT. 1957. The ctriated band (Lepidoptera 
chiefly Arctiidae). Lepid. News 11: 147-150. 

Haske, P. T. & P. Betron. 1956. Electrical responses of certain lepidopterous 
tympanal organs. Nature 177: 139-140. 

Howven, H. F. & L. E. C. Line. 1973. Scanning electron microscopy: low 
magnification pictures of uncoated zoological specimens. Science 179: 386- 

Roeper, K. D. 1962. The behaviour of free flying moths in the presence of 
artificial ultrasonic pulses. Anim. Behav. 10: 300-304. 

1967a. Turning tendency of moths exposed to ultrasound while in sta- 

tionary flight. J. Insect Physiol. 13: 873-888. 

1967b. Nerve Cells and Insect Behavior. Harvard, Cambridge. revised 


. & A. E. Treat. 1957. Ultrasonic reception by the tympanic organ of 
noctuid moths. J. Exp. Zool. 134: 127-157. 

Roruscumtp, M. 1961. Defensive odour and Mullerian mimicry among insects. 
Trans. Roy. Entomol. Soc. London 113: 101-121. 

1965. The stridulation of the garden tiger moth. Proc. Roy. Entomol. 

Soc. London (C) 30: 3. 

. & R. T. Aupry. 1971. Toxins in tiger moths (Arctiidae: Lepidoptera), in 
Pesticide Chemistry, A. S. Tahori, ed., Gordon and Brech, London, p. 177-182. 

SCHALLER, F. & C. Tmwm. 1950. Das Horvermogen der Nachtschmeterlinge. Z. 
Vergl. Physiol. 32: 468-481. 

TreEAT, A. E. 1955. The response to sound in certain Lepidoptera. Ann. Entomol. 
Soc. Amer. 48: 272-284. 


On 22 July 1973, while collecting Lycaeides melissa samuelis (Edwards) in the 
vicinity of Griffith, Lake Co., Indiana, I took a pair of Problema byssus (Edwards), 
a slightly worm male and a fresh female, the first recorded from that state that I 
had knowledge of at the time. Identification was kindly verified by Mr. Ernest M. 
Shull, co-author of an annotated list of the butterflies of Indiana (1972, J. Lep. 
Soc. 26: 13-24). According to Mr. Shull who, along with Mr. F. Sidney Badger, 
has carried out intensive collecting and study in Indiana, these are the first of- 
ficially recorded specimens for that state. Both specimens have been placed in the 
private collection of Mr. Shull. 

On 24 June 1973, I took two fresh males of the color form pallida of Thymelicus 
lineola (Ochsenheimer) in Spears Woods Forest Preserve, Cook Co., Illinois. Ac- 
cording to word received from the Illinois Natural History Survey, these are the 
first records from Illinois. Both specimens have been placed in the permanent 
Survey collections. 

IRWIN LeEuw, 1219 Crystal Lake Road, Cary, Illinois 60013. 



Department of Biology, Towson State College, Baltimore, Maryland 21204 


Department of Entomology, University of Illinois, Urbana, Illinois 61801 

During the course of our studies of the cecropia moth, Hyalophora 
cecropia (L.), we found a large proportion of cocoons in low shrubs. 
We suspected that many of these shrubs, particularly species of Juniperus 
and Taxus, do not support larval growth, and that the presence of cocoons 
on them is evidence that pre-spinning larvae wandered to them from the 
foodplant. Most of the shrubs in question are not included on published 
lists of the hostplants of cecropia, but since this does not prove that 
cecropia larvae could not feed on them, we made the feeding trials 
described below. 

There is some doubt as to whether or not all of the species on the 
published “foodplant lists” of cecropia are actually eaten by cecropia 
larvae. Brodie’s (1882) list is reliable since he included only plants 
on which larvae had been found feeding in the field. On the other 
hand, Marsh’s (1937) list is of dubious value since he included plants 
on which he had found cocoons but had not seen larvae. Tietz (1958) 
compiled a long list, but, unfortunately, did not state the evidence upon 
which the plants were included. 


Most cocoons were collected in residential areas, principally in the 
twin cities of Champaign and Urbana, Illinois, although a few were 
found in nearby small towns. A small number came from rural areas, 
mostly from ditch banks, railroad rights-of-way, roadsides and fence rows. 

The larvae used in feeding trials were the progeny of wild parents 
which had been collected as described above. After mating in the 
laboratory, females were placed in large paper bags where they ovi- 
posited. Before hatching occurred, small bits of paper bearing the eggs 
were snipped out and transferred to petri dishes. The unfed first instars 
used in the first series of tests were indiscriminately selected within a 


half-hour of hatching. The partly grown larvae used in the second 
series were indiscriminately selected from groups of larvae reared in 
nylon mesh sleeves on apple trees (Malus pumila) essentially as described 
by Telfer (1967). 

Plants of 118 species were tested for their ability to support the growth 
of first instar larvae. Each species was tested with at least three replicates. 
Species on which no larvae survived the first stadium in the initial test 
were retested with an additional three replicates. Each replicate con- 
sisted of ten larvae confined with foliage in a 10 cm petri dish lined 
with a disc of moist filter paper. The newly hatched larvae were weighed 
in groups of ten and immediately placed in the dishes. Undamaged 
foliage, collected daily from plants growing in full sunlight, was sealed 
in plastic bags and stored in a refrigerator until used later the same day. 
The dishes were kept under constant illumination and at a temperature 
of 23 = 1° C. At least once a day fresh food was added, and dead 
larvae, left over food, and feces were removed. Feces were dried im- 
mediately, and eventually the aggregate for each replicate was redried 
to a constant weight at 100° C (see Waldbauer, 1964). This weight was 
divided by the sum of the number of larvae feeding on each test day 
to yield the mean dry weight of feces passed per larva per day. Larvae 
were weighed and considered to have survived the stadium as soon as 
they had spun a molting pad. Dishes with moist filter paper but no 
food, and dishes with Acer saccharinum foliage served as negative and 
positive controls respectively. A group of tests was started on each of 
three days. Each group had its own controls, but since the controls 
differed by very little they have been lumped in Table 2. 

A few plants were also tested with two groups of partly grown larvae. 
Each group consisted of five larvae confined in a sleeve on a branch 
of a living plant in the field. They were transferred to the test plant 
from apple foliage, one group just before the molt to the fifth stadium 
and the other on the seventh day of the fifth stadium. Apple branches 
with leaves served as positive controls, and defoliated branches of the 
test plant as negative controls. 


Table 1 lists the plants on which we found 1% or more of the cocoons 
collected during this study. Almost all of the cocoons from rural areas 
were on Salix interior, a shrubby willow which grows wild on ditch 
banks and in other moist places, but is not planted in urban areas. A 
few of the other listed species grow wild in this area, but with few 


TABLE 1. The location of cecropia cocoons found during the collecting seasons 
of 1967-68, 1968-69 and 1969-70. Only those plants on which 1% or more of 
the total was found are named. Species which do not serve as foodplants for 
cecropia are marked with an asterisk. 

Cocoons Found 

Locations Number % of total 
Acer saccharinum 1. OTA 34.8 
*Juniperus spp. 278 9.0 
Salix interior Rowlee 254 8.2 
Betula pendula Roth. 168 5.4 
Acer rubrum L. 136 4.4 
Rhamnus frangula 1. 132+ 4.3 
Malus spp. (Including pumila) D7 A.J 
*Ligustrum vulgare L. 115 Ben 
*Taxus media Rehd. 102 oo 
Betula populifolia Marsh. 90 2.9 
Platanus occidentalis L. 66 Del 
Betula papyrifera Marsh. 52 ihe 
Cornus stolonifera Michx. and 
G. alba. 46+ 185 
*Euonymus spp. (not alatus) 38 1e?, 
Other plants 300 1's 
Fences and buildings Syl ie 
3,084 99.8 

+ See text. 

exceptions we found cocoons only on cultivated specimens in urban 

The data of Table 1 do not reflect the importance to cecropia in this 
area of Rhamnus frangula and the two species of Cornus. In an earlier 
study (Waldbauer & Sternburg, 1967) we found cocoons abundantly 
on Cornus stolonifera and C. alba. However, during the present study 
we did little collecting from Cornus because most plants of these species 
in this area were included in another study of cecropia. R. frangula was 
formerly scarce in this area, but has become a popular hedge plant 
since we began our studies of cecropia in 1965; the great majority of 
cocoons from R. frangula were collected during the last year of the 
present study. 

Over 12% of the cocoons were found on shrubby conifers, Juniperus 
spp. and Taxus media, but only if these conifers were close to trees, 
particularly Acer saccharinum and Betula spp., which are important 
foodplants of cecropia in this area. We hypothesized that the larvae do 
not feed on these conifers, but migrate to them from their foodplants 
when they are ready to spin cocoons. The observations recorded below 


TaBLE 2. Survival, growth and feces production of first instar cecropia larvae 
on the leaves of various woody plants. All plants were tested with three replicates 
of ten larvae each except those marked with an asterisk, which were tested with 
six replicates of ten each, and A. saccharinum and moist filter paper which were 
tested with 9 replicates of ten each. 

Duration of Mg fresh Mg dry feces/ 

instar (days) weight gained larva/day 
Plants eee Mean Range Mean Range Mean Range 
Plants listed in Table 1 

Acer rubrum LL. S616 610) 22610-6107 224 (2408 39%) 379-410 
A. saccharinum L. O50 63 5.02/00 202 le7Wse BG SO Ssteys) 
Betula papyrifera Marsh. 83.3 3.8 3.5-4.0 329 309-345 39.9 34.1-44.2 
B. pendula Roth. V0.0 46 4s) eet Sees) SiG WME exay 
Cornus alba L. CRS 4:3 d@s50- B85 IIGBEWES IG Ue fe tkeyy 
C. stolonifera Michx. Oe AiG AOS Ws ONG LOs) DUKE a Is) 
Euonymus alatus Sieb. 900 88 8.0-9.5 180 166-203 22.7 20.4-24.3 
E.: fortunei Trucz.* 0 — - 0.1 0.1-0.3 
E. yedoensis Koeh.* 0 — - 29  1.9-49 
Juniperus chinensis L.* AG xeh0) = 80 = 46 2.9-6.8 
J. communis L.* 0 = = O83 OvleOe 
J. procumbens Endl.* 0 - - 49 2.0-8.9 
J. sabina \.* 0 — — 0.7 0.5-0.9 
J. virginiana L.* 0 — - 0.3 0.2-0.3 
Ligustrum vulgare L. a) dl) - 244 — IAQ) PMN5G 
Malus adstringens Zabel. 96.6 5.3 5.0-6.0 279 232-264 30.9 26.5-33.3 
M. arnoldiana Sarg. WS.)  — H.83 D055 SIL SOs sO) BIL IL Ae} 5858}. 
M. atrosanguinea Scheid. 96.6 5.3 5.0-6.0 252 232-264 30.8 28.0-32.9 
M. floribunda Sieb. Co. 6.0. BOQ Ise ISSO wr) VES 4 
M. pumila Mill. OBO 4 AO.0 Bibl SOE wl 2ee  aulsr sg! 

occidentalis L. 100.0 BS BSS SEO ONPG Gey Wes) 
Rhamnus frangula L. 100.0 43 404.5 288 282-300 27.2 25.8-29.8 
Salix interior Rowlee C66 B5 Bis — Diy Waals Boxe} sO Casts 
Taxus media Rehd.* 0 = - OG 0207 

Larix decidua Mill. S00 BO “5O5.0 We) esas) Viskish) WS Oakes 
L. laricina (Du Roi) 

K. Koch Srey | ALS) 7 eae 5 Ole Z265—oC amo le oO Aol 
Pinus flexilis James 0 — — 2.6 1.0-5.8 
P. nigra Arnold* 0 ~ — Ol OL1=0:2 
P. strobus L. EGE See nOSeo Ss ie eIO=17 2 6) 3:6—1180 
P. sylvestris 1.* 0 — _ 0.5 0.40.8 
Taxodium distichum 

(its) wich: SOO) — PAIL — 1.8 0.5-3.7 
Thuja occidentalis L.* 0 = = Oa ED =209 
Tsuga canadensis 

(les) Carn. S00 6 COs IDA “Sabi IRRy eZ H0) 
Moist filter paper 0 — = OP O4ep 4 


show this hypothesis to be correct and, furthermore, show that the cocoons 
spun on certain angiosperms (see Table 1) are probably also spun by 
migrants from other plants. 

We have found wild third to fifth instar larvae in the field feeding 
on the following plants: Acer platanoides, A. saccharinum; Betula 
papyrifera, B. pendula, B. populifolia; Cornus alba, C. stolonifera; 
Lonicera fragrantissima Linden and Paxton, L. tartarica L.; Malus pumila, 
Malus spp.; Paeonia officinalis L.; Prunus serotina; Rhamnus frangula; 
Rhus typhina; Salix interior; and Spirea alba. We have observed success- 
ful development to the adult stage on all of these except A. platanoides, 
B. populifolia, Malus spp. other than M. pumila, and S. alba. We have 
no reason to suspect that the latter would not support complete develop- 

Table 2 gives the results of some of the feeding trials with first instars. 
In the upper section of the table appear the plants on which most 
cocoons were found (see Table 1). In the lower section are listed other 
conifers, some of which, to our surprise, supported good growth. 

Fifteen of the species on which we had found large numbers of 
cocoons supported apparently normal growth to the end of the first 
stadium; 83% or more of the test larvae survived the stadium, gained a 
fresh weight of 180 mg or more—usually in six days or less, and ate at 
a more or less normal rate as judged by the rate at which they passed 
feces. This does not prove that development could have been completed 
on these plants. However, we have found fifth instars feeding in the 
field on all of these plants except Acer rubrum and Platanus occidentalis. 
Thus there is almost no question that cecropia can complete development 
on thirteen of these plants, and there is no reason to assume that it cannot 
complete development on A. rubrum and P. occidentalis. It is, therefore, 
unlikely that cocoons found on these plants were spun by larvae which 
had wandered from some other species of plant. 

The other plants on which large numbers of cocoons had been found, 
nine species of Ewonymus, Juniperus, Ligustrum and Taxus, were usually 
eaten only in minute quantities and did not support growth or, at best, 
supported the slow growth of only one larva out of a group of 30 or 
60 (see Table 2, J. chinensis and L. vulgare). The inability of these 
plants to support the growth of first instars proves that they are not 
usual food plants of cecropia. Therefore, cocoons found on them must 
have been spun by larvae which had moved from some other species 
of plant, the movement probably occurring only after growth had been 

It is possible that plants which do not support first instars may sup- 




TABLE 3. Survival of partly grown cecropia larvae transferred from Malus pumila 
to other plants. Each plant was tested with two groups of five larvae. 

Larvae transferred at: 

end of 4th instar 7th day of 5th instar 
No. surviving to: No. surviving to: 
Plants pupa adult pupa adult 
Test plants 
Juniperus chinensis L. 0 0 0 0 
J. communis L. 0 0) 0 0) 
J. procumbens Endl. 0 0 0 0 
J. virginiana L. 0 0) 0 0 
Ligustrum vulgare L. 4 1 3 0 
Taxus media Rehd. 0 0 0 0 
Malus pumila Mill. 5 5 5 5 
Defoliated branch 0 0 0 0 

port the growth of partly grown larvae which may move to them 
from a more favorable plant. We have seen no evidence of such moves, 
and doubt that they are of more than rare occurrence. Furthermore, 
feeding tests with partly grown larvae (Table 3) indicate that although 
an occasional partly grown migrant might survive by eating the foliage 
of Ligustrum vulgare, there will be no survival on Juniperus or Taxus, 
even if the migrants feed on one of the usual foodplants until the middle 
of the last stadium. Taxus is probably toxic since larvae which ate small 
quantities died sooner than controls on a defoliated branch of Taxus. 

Three of the four species in the genus Hyalophora apparently feed 
only on non-conifers, while the remaining species, the northern H. 
columbia (S. I. Smith), appears to feed exclusively on the tamarack, 
Larix laricina, (Ferguson, 1971). It is thus of more than passing interest 
that some conifers, most notably Larix laricina, L. decidua, and Tsuga 
canadensis, support good survival and growth by cecropia during the 
first stadium (Table 2). Furthermore, during the summer of 1973 we 
transferred five young third-instar cecropia from an apple to a small 
European larch (L. decidua) growing outdoors. Two disappeared from 
the sleeve, but the remaining three spun apparently normal cocoons. 
Whether or not they survive to emerge as adults remains to be seen 
since they are still in diapause. Recently Collins (1973) reported that 
the two western species can survive on some conifers. H. gloveri 
(Strecker) has been reared on L. decidua, L. laricina and Pseudotsuga 
menziesii (Mirb.). H. euryalus (Boisduval) has been found in the field 
feeding on P. menziesii and has been reared in captivity on the same 


plant. It would seem that all Hyalophora are potentially able to utilize 
conifers, but that this ability has been selected for only in H. columbia. 

Many plants in addition to those listed in Table 2 were tested with 
first instars, but it would require too much space to list the data in 
detail. Plants on which over 70% of the larvae survived the first stadium 

Acer platanoides Michx., A. negundo L.; Aesculus octandra Marsh.; Carya ovata 
(Mill.) K. Koch, C. illinoensis (Wangenh.) K. Koch; Castanea mollisima Bl.; 
Chaenomeles lagenaria Koidz.; Cornus florida L., C. racemosa Lam., C. sanguinea 
L.; Cotoneaster multiflora Bge.; Crataegus crus-gali L., C. molli Scheel.; Diospyros 
virginiana L.; Fraxinus pennsylvanica Marsh.; Hamamelis virginiana L.; Juglans 
nigra L.; Liquidambar styraciflua L.; Lonicera tartarica L.; Maclura pomifera (Raf. ) 
Schneid.; Nyssa sylvatica Marsh.; Populus deltoides Bartr., P. laurifolia Ledeb.; 
Prunus americana Marsh., P. serotina Ehrh.; Pyrus communis L.; Quercus alba L., 
QO. imbricaria Michx., Q. macrocarpa Michx., Q. muhlenbergii Engelm., Q. rubra 
L.; Rhus typhina L.; Robinia pseudoacacia L.; Salix babylonica L.; Sambucus 
canadensis L.; *Sanicula smallii Bicknell; Spirea alba Du Roi; Syringa chinensis 
Willd.; Tilia americana L., T. euchlora Koch., T. tomentosa Moench.; Viburnum 
dentatum L., V. tomentosum Thunb. 

Plants on which there was survival, but less than 70% are: 

Acer saccharum Marsh.; *Asclepias syriaca L.; Betula alleghaniensis Britton, 
B. nigra L.; Cotinus americana Nutt.; Deutzia lemoine Hort.; Elaeagnus angustifolia 
L.; Forsythia viridissima Lindl.; Fraxinus americana L.; Ginko biloba 1.; Gleditsia 
triacanthos L.; Halesia carolina L.; *Lactuca scariola L.; Lonicera japonica Thunb.; 
Morus alba L.; *Plantago rugelii L.; Rhus glabra L.; Salix nigra Marsh.; Syringa 
vulgaris L.; *Taraxacum officinale Weber; Tilia platyphyllos Scop.; Ulmus americana 
L., U. carpinifolia Gleditsch., U. parvifolia Jacq. 

Plants on which no larvae survived are: 

*Adiantum pedatum L.; Ailanthus altissima (Mill.) Swingle; Asimina triloba 
Dunal.; Campsis radicans (L.) Seem.; Catalpa bignonioides Walt.; Celtis occidentalis 
L.; Cercis canadensis L.; *Chenopodium album L.; Kolkwitzia amabilis Graebn.; 
Liriodendron tulipifera L.; *Nicotiana tabacum L.; Philadelphus coronarius L.; 
Populus alba L.; *Sonchus asper (L.) Hill; *Verbascum thapsus L.; *Viola sp.; 
Vitis sp.; *Zea mays L. 

In the above list herbaceous plants (as opposed to woody) are marked 
with an asterisk. Larvae survived on only five of the eleven non-woody 
plants tested. These were: Asclepias syriaca—30%, Lactuca scariola— 
3%, Plantago rugelti—53%, Sanicula smallii—83% and Taraxacum of- 
ficinale—63%. Almost without exception the recorded natural food- 
plants of cecropia are woody; the only indisputable exception is Paeonia 
officinalis (see above, and Waldbauer & Sternburg, 1967). 

Table 2 shows considerable variation in response to the various species 
of acceptable plants, particularly in the duration of the instar, weight 
gained and the weight of feces passed per day. Large differences in 
the latter value (cf., for example, Malus pumila and Cornus alba) sug- 
gest large differences in the rates at which different plants are eaten. 

VOLUME 28, NUMBER 3 219 

However, the weight of feces is only an approximate indication of the 
rate of intake since the former will vary not only with the weight of 
food ingested, but also with the proportion of the ingested food which 
is assimilated and expended for growth and the maintenance of metabo- 
lism (Waldbauer, 1964). 

This study makes the following major points: 

1. It confirms past observations that cecropia larvae are able to feed 
and survive on a wide variety of woody angiosperms. 

2. It shows that neither first nor fifth instar larvae are able to survive 
on certain plants on which cocoons are commonly found, establishing 
that the larvae must have moved to them after completing their feeding 
on some other species of plant. 

3. It shows that first instars are able to survive on certain species 
of conifers, including Larix laricina, the foodplant of the closely related 
Hyalophora columbia. 


Bropiz, W. 1882. Food plants of Platysamia cecropia. Papilio 2: 32-33. 

Coxuins, M. M. 1973. Notes on the taxonomic status of Hyalophora columbia 
(Saturniidae). J. Lepid. Soc. 27:225-235. 

FERGUSON, D. C. in Dominick, R. B. et al. 1972. The Moths of America North 
of Mexico, Fascicle 20.2B, Bombycoidea (in part). Classey, London. p. 155- 

Marsu, F. L. 1937. Ecological observations upon the enemies of Cecropia, with 
particular reference to its hymenopterous parasites. Ecology 18: 106-112. 

TELFER, W. H. 1967. Cecropia, in F. H. Wilt and N. K. Wessels, eds., Methods 
in Developmental Biology. p. 173-182. Crowell, New York. 

Tretz, H. M. 1959 (?). The Lepidoptera of Pennsylvania, a Manual. Pennsylvania 
State Coll. Sch. Agr. & Agr. Exp. Sta. 194 p. 

WALDBAUER, G. P. 1964. Quantitative relationships between the numbers of fecal 
pellets, fecal weights and the weight of food eaten by tobacco hornworms, 
Protoparce sexta (Johan.) (Lepidoptera: Sphingidae). Entomol. Exp. & Appl. 

. & J. G. SterNBuRG. 1967. Host plants and the locations of the baggy and 

compact cocoons of Hyalophora cecropia (Lepidoptera: Saturniidae). Ann. 

Entomol. Soc. Amer. 60: 97-101. 

Note AppED IN Proor: The three pupae from larvae matured on Larix decidua 
produced three normal adult moths in 1974: 21 May, 4; 29 June, 6; 30 June, @. 




213 Mt. Salus Drive, Clinton, Mississippi 39056 

Variation in size of individuals of Euptoieta claudia (Cramer) has been 
reported, but no quantitative data have been found in the literature. 
Mather & Mather (1958) wrote of Mississippi that, “February specimens 
are characteristically very small.” Harris (1972) wrote of Georgia that, 
“The individuals of E. claudia vary in size, and an interesting series may 

1 Contribution No. 271, Bureau of Entomology, Division of Plant Industry, Florida Department 
of Agriculture and Consumer Services, Gainesville, Florida 32601. 

2 Research Associate, Florida State Collection of Arthropods, Division of Plant Industry, 
Florida Department of Agriculture and Consumer Services, Gainesville. 

10 38 MALES ) 
9 4 
8 5 
7 6 
6 7 
N 2 Z 
4 9 
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2 11 


—“Nwhl_ unan ovo o 

20 (22 24 26 28 30 32 34 36 38340eeeZ 


Fig. 1. Forewing-length distribution of 75 Mississippi specimens of Euptoieta 


NW eR 



Dw ee 
S&S) ce) ©) 




TABLE 1. Data on 75 specimens of Euptoieta claudia from Mississippi. 


Feb 52 
Feb 49 
Feb 49 


FW length 

Locality Collector (mm ) Sex 
Clinton, Hinds B. Mather 21 3 
Clinton, Hinds B. Mather 20 3 
Clinton, Hinds B. Mather 20 3g 
Brooklyn, Forrest B. D. Valentine 26 J 
Clinton, Hinds M. & E. Roshore 26 3 
Jackson, Hinds B. Mather Dil J 
Vicksburg, Warren B. Mather 33 2 
Vicksburg, Warren B. Mather 26 3 
Clinton, Hinds B. Mather Sil Q 
Jackson, Hinds B. Mather 29 3 
Clinton, Hinds B. Mather Dil 3} 
Jackson, Hinds B. Mather 28 3 
Bynum Mounds, Chickasaw M. & E. Roshore 29 3 
Thrasher, Prentiss M. & E. Roshore oo Q 
Brownsville, Hinds M. & E. Roshore 34 Q 
Gulfport, Harrison B. Mather 28 2 
Bay St. Louis, Hancock B. Mather 28 3 
Clinton, Hinds M. & E. Roshore 29 & 
Clinton, Hinds M. & E. Roshore Bw A 
Clinton, Hinds M. & E. Roshore 29 A 
Clinton, Hinds M. & E. Roshore 30 3 
Clinton, Hinds B. Mather 32 Q 
Clinton, Hinds B. Mather 35 Q 
Clinton, Hinds B. Mather 34 Q 
Clinton, Hinds B. Mather Di & 
Clinton, Hinds B. Mather 29 3 
Clinton, Hinds B. Mather 36 Q 
Jackson, Hinds B. Mather 35 Q 
Clinton, Hinds B. Mather 36 Q 
Brownsville, Hinds B. Mather 35 Q 
Clinton, Hinds B. Mather 26 3 
Jackson, Hinds B. Mather 34 Q 
Clinton, Hinds B. Mather 3¢ Q 
Clinton, Hinds B. Mather 32 Q 
Clinton, Hinds M. & E. Roshore 35 fe) 
Tishomingo St. Pk., 

Tishomingo M. & E. Roshore 32 2 
Bovina, Warren B. Mather 33 Q 
Jackson, Hinds B. Mather 28 3 
Jackson, Hinds B. Mather 28 3 
Clinton, Hinds B. Mather 35 2 
Clinton, Hinds B. Mather 37 2 
Clinton, Hinds B. Mather 36 2 
Brownsville, Hinds B. Mather Be Q 
Clinton, Hinds B. Mather 32 3 
Bolton, Hinds B. Mather 30 3 
Clinton, Hinds M. & E. Roshore 36 2 



TABLE 1. (Continued) 

FW length 

Date Locality Collector (mm ) Sex 
2 Sep 56 Brownsville, Hinds B. Mather 29 3 
2 Sep 57 Bolton, Hinds M. & E. Roshore 31 3 
5 Sep 49 Clinton, Hinds B. Mather 29 3 
12 Sep 71 Clinton, Hinds B. Mather 35 2 
19 Sep 53 Waynesboro, Wayne B. Mather 29 a 
23 Sep 72 Jackson, Hinds B. Mather 35 2 
25 Sep 54 Jackson, Hinds B. Mather 34 2 
26 Sep 68 Hattiesburg, Forrest E. Reid 36 2 
26 Sep 68 Hattiesburg, Forrest E. Reid 27 3 
26 Sep 68 Hattiesburg, Forrest E. Reid 27 3 
28 Sep 58 Jackson, Hinds B. Mather 35 2 
3 Oct 59 Clinton, Hinds M. & E. Roshore 31 2 
sy (Oleh SP Clinton, Hinds B. Mather 38 Q 
5 Oct 58 Brownsville, Hinds M. & E. Roshore 28 3 
6 Oct 51 Clinton, Hinds B. Mather 29 a 
6 Oct 51 Clinton, Hinds B. Mather 33 Q 
97Ock bil Ridgeland, Madison B. Mather 29 3 
11 Oct 69 Ft. Adams, Wilkinson B. Mather Bz Q 
12 Oct 52 Clinton, Hinds B. Mather 28 3 
2 Ock a2 Clinton, Hinds B. Mather reill 3 
12 ‘Oct 69 Pinckneyville, Wilkinson B. Mather Or: 3 
12 Oct 69 Pinckneyville, Wilkinson B. Mather og Q 
13 Oct 56 Vicksburg, Warren B. Mather a2 Q 
13 Ockn 56 Jackson, Hinds B. Mather 3o Q 
l5rOck 55 Jackson, Hinds B. Mather 30 3 
250Ock 71 Jackson, Hinds B. Mather 33 2 
4 Nov 51 Clinton, Hinds B. Mather ot Q 
7 Nov 48 Clinton, Hinds B. Mather 25 3} 
ITSNovibk Hermanville, Claiborne B. Mather 23 3 

be obtained. Very small specimens may be found at times, especially 
in early spring, and unusually large ones may be found in the summer.” 
Rahn (1972) reported on five specimens taken 18-21 August 1970 in 
North and South Dakota. He stated that these showed a “wing span 
range from 1%” to 2”.”. Measurements made on the figure in his paper, 
adjusted for scale reduction in reproduction, indicate forewing lengths 
of 20, 24, 30, 30, and 32 mm. 

A group of 75 Mississippi specimens was examined; results are given in 
Table 1. Forewing-length distributions for males and females are shown 
in Fig. 1. As previously noted, very small specimens have been taken 
in February. Other small specimens were taken in November; but there 
does not appear to be the progressive increase in size through the season 
that characterises such species as Colias eurytheme Boisduval or Papilio 
glaucus Linnaeus. The major factors affecting size in E. claudia appear 



to be (a) sex, females are typically significantly larger than males; and 
(b) winter, November and February specimens are characteristically 
smaller than average. The size range of the Mississippi series is 20 to 
38 mm (1 to 1.9). The range of the series of five August specimens 
reported by Rahn (1972) was 20 to 32 mm (1 to 1.6). The size range 
for six Mississippi August specimens was 28 to 37 mm (1 to 1.3). 


Harris, L., Jr. 1972. Butterflies of Georgia. Univ. Okla. Press, Norman. p. 280. 

Martner, B. & K. MAtHer. 1958. The butterflies of Mississippi. Tulane Stud. Zool. 
6: 63-109. 

Raun, R. A. 1972. A dwarf form of Euptoieta claudia (Lepidoptera: Nymphalidae). 
Great Lakes Entomol. 5: 101. (Reproduced, with minor editorial changes 
and reversal of the figure, in J. Res. Lepid. 11: 174.) 


In Jones’ “Annotated Check List of the Macrolepidoptera of British Columbia” 
(1951), Phyciodes mylitta is listed as occurring in a number of localities in mainland 
British Columbia including the coast adjacent to Vancouver Island, but there were 
at the time no records for Vancouver Island. I have myself collected intensively 
over a large part of Vancouver Island during the past 30 years, without encountering 
P. mylitta before 1972. 

In August of that year P. mylitta tumed up in Bright Angel Park near Duncan. 
The first specimens were handed to me by a friend, Mrs. Betty McKinnon, who 
said that they were then quite plentiful in the park area. Looking at the butterflies 
through the semi-transparent envelopes in which they were packed, I took them 
for Phyciodes campestris Behr. which I have often taken on Vancouver Island, though 
I was surprised at the late date, 10 August. I did not examine them more closely 
until late the following winter, when I at once saw that they were not P. campestris. 
Later, I sent some examples to Dr. dos Passos, who pronounced them typical 
Phyciodes mylitta. 

The following April, P. mylitta showed up in a number of localities from Victoria 
on the southern tip of the Island north about 40 miles to Chemainus. Either the 
species had maintained itself in such small numbers as to escape detection, and 
then suddenly exploded; or it had gained access to Vancouver Island two years ago 
and built up a large population with amazing speed. In some respects the case 
resembles that of Coenonympha tullia, which after being confined to the immediate 
vicinity of Victoria until about 1965, suddenly started to spread northward, again 
coming to a stop near Chemainus. The latter species, however, was always very 
common near Victoria. 

I cannot account for the flight season for P. mylitta as given by Jones, he states 
merely “June.” Last year I observed a spring brood starting to fly very early in 
April, followed by a summer flight in July and August. P. campestris here is 
single brooded, flying in June and early July. 

Ricuarp Guppy, Thetis Island, British Columbia, Canada. 



101 Avenida Norte #322, San Salvador, El Salvador 


Escuela Nacional de Agricultura, San Andres, Depto. La Libertad, El Salvador 

Nocturnal gregarious roosting is known to exist in a variety of reputedly 
unpalatable species of butterflies (Carpenter, 1931; Crane, 1955; Jones, 
1930; McFarland, 1970; McNeil, 1937; Poulton, 1931; and our personal 
observations on the phenomenon in adults of Dryas iulia iulia ( Fabricius ), 
Heliconius petiveranus Doubleday and H. charitonius L.). According 
to A. M. Young (pers. comm.), one case of nocturnal gregarious roosting 
of a nymphalid, Marpesia bernia (Hewitson), has been studied. But 
apparently a semi-permanent or seasonal, 24 hours a day, six months 
a year kind of roosting has not been known to exist in any kind of 

As stated in our preliminary report on communal resting of Smyrna 
karwinskii (1973), we have observed, annually since 1962, adults of 
Smyrna karwinskii (Geyer) resting during the day in groups ranging 
from 10 to more than 100 individuals, of mixed sex ratio, in cavities of 
lava walls and tree trunks, and on the underside of concrete slabs 
roofing alleys between cabins at Cerro Verde (a mountain of about 
2000 m elevation overlooking the WNW slope of the Izalco Volcano, 
about 50 km from San Salvador). In total we have observed the 
phenomenon 37 days during the months of January, February, March, 
April, August, November and December in the course of eleven years. 
In each of these months, in different years, the observations have been 
repeated several times, except in August (only one time during 1965). 
In all instances the roosting aggregations observed were situated in the 
shade, away from direct sunlight. The locations chosen by the groups 
were always protected from the northern winds, which are usually 
quite strong from December through February, principally on mountain 

On 31 March 1972 we observed five roosting aggregations very closely. 
One contained 15 adults, a second 20, two about 50 each and a fifth more 

VoLUME 28, NUMBER 3 225 

than one hundred. Monthly trips were made to the same place from 
June to November, but not a single group was found during that period. 
On 23 December, two groups were located, one consisting of 123 
individuals (Fig. 1), and a second, some three meters from the first, 
of 39 (Fig. 2). As usual, both males and females were present, but no 
sexual activity was noticed. From time to time individuals would depart 
from the groups, presumably in search of food resources, and sporadically 
individuals would join the groups. 

In order to determine the sex ratio in the aggregations, the one that 
contained 123 individuals was captured early in the morning of 23 
December 1972, using a big bag made of stiff plastic sheet, that when 
flattened acted as a big paper envelope, immobilizing the butterflies 
without damaging them. To cause the butterflies to move into the bag, 
the mouth of it was applied to the concrete, enclosing the whole group, 
and then moved back and forth until all individuals, except 15 that 
escaped, were inside. The bag was flattened, keeping the butterflies 
motionless on their sides, and one by one they were extracted, quickly 
sexed and counted. Sexing this species is easy due to their sexual 
dimorphism. The total was 42 males and 66 females. As they were 
freed, they darted to the surrounding wood, alighting on tree trunks 
and rocks. During the following 5 hours, there was much flying activity: 
males chasing females and other males. Some individuals started to 
alight singly under the roofing slabs, scattered over an area of roughly 
40 m. By the time we left we counted 38 individuals. None joined the 
smaller group, nor had they started to form a new aggregation. 

We tried to mark the second smaller group with red spray from 
a distance, but the group started to disintegrate when the mist reached 
it, so we stopped this method. Out of the original 39 butterflies in the 
group, only 18 stayed. We tried then to mark the remaining butterflies 
individually by capturing each one by hand, giving it a light spray 
on the right rear underwing and putting it back in place, but the first 
individual so treated did not stay with the group and flew away. The 
rest were then left alone for future observations. 

On 28 January 1973, about a month later, we visited the same place. 
The large group was not found. The smaller, though, was at the same 
place it had been on 23 December, and it had about the same number 
of individuals as when first sighted. Even the marked one was there, 
standing conspicuously in the second row. Unfortunately on the fol- 
lowing trip to the locality this group was not found again, having been 
disturbed the week before by some curious tourists, according to the 
report of one of the guards. 


On various occasions females were captured from aggregations and 
dissected. None had eggs in the abdomen, but considerable amounts 
of fat tissue were present. 

It has been noticed that during the wet season the species is found 
in lower lands, where the foodplants (several species of Urticaceae), 
are very abundant. During the wet season it is rare to find adults of 
Smyrna karwinskii wp in Cerro Verde. Not so during the dry season. 

We have collected eggs and larvae of this species during the wet 
season in the same habitat we have collected eggs and larvae of its 
more common close relative Smyrna blomfildia datis Friuhstorfer, which 
is seldom found at high altitudes (the highest record for S. b. datis in 
El Salvador is one adult captured on the slopes of Cerro Verde, ca. 1600 
m, by S. R. Steinhauser in October 1967, pers. comm.), yet we have 
never found eggs or larvae of S. karwinskii in Cerro Verde. It is to be 
noted that the larvae of S. karwinskii are easily distinguished from the 
larvae of S. blomfildia datis by their color. S. karwinskti have the body 
and spines brown, while S. blomfildia datis presents four different 
morphs: body and spines greenish white, body greenish white with 
black spines, body mostly black with light spines and body mostly black 
with black spines. In shape and head the two species are much alike. 


Since the phenomenon was first observed, back in 1962, it had been 
our assumption that the butterflies were grouped at places where water 
filtrated and that they were drinking there. But on 31 March 1972 we 
had a chance to observe five groups from a short distance for a period 
of seven hours. Not once did we see any individual uncoil its proboscis, 
nor did we notice any moisture in the concrete, thus eliminating that, 
as well as the alternative explanation that the grouping was formed to 
allow use of each other's excreta to recycle fluids, as has been reported 
by Hessel (1966) for single individuals of Agathymus aryxna (Dyar). 
The fact that the groups are always formed in the shade, away from 
direct sunlight, also eliminates the possibility that receiving solar heat 
plays a role in causing the aggregations. 

In the unpalatable species, gregarious nocturnal roosting has been 
viewed as an evolutionary behavior acquired to enhance their unpleasant 
scent and therefore the chances to effectively deter any approaching 
predator. Smyrna karwinskii adults are not reputed to be distasteful 
to predators, but no experiment that we know of has been carried out 
in this respect. We have observed that the larvae when molested extrude 
a gland located anterad of the prothoracic legs, as its relatives S. 

VoLUME 28, NUMBER 3 Pag 

blomfildia datis, Colobura dirce LL. and Historis odius Fabricius do, 
emitting a scent very faint to humans, presumably to repel potential 
vertebrate predators (Hemiptera have been found feeding on Smyrna 
spp. larvae). This makes us doubt the palatability of the adults. If 
after appropriate experiments, this species proves to be distasteful to 
such predators, the communal roosting habits could be explained for 
the reasons given above. This mechanism would act only as a chemical 
repellent, having a passive role, as it is evident that the individuals in 
the congregation do not have the ability to communicate to the other 
members of the group when danger is imminent, as individuals can 
be captured by hand from any place in the group and at any time 
of the day without causing a reaction from the rest of the individuals. It is 
necessary to be rather rough to obtain a mass response from the whole 
aggregation. When this is done the individuals disperse in all directions 
producing an audible rustling noise with the wings, somewhat like 

Another possible benefit that the congregated individuals seem to 
derive from their communal roosting is the mimetic effect obtained: 
the groups look like a dried moss or lichen formation, at least to humans. 

The fact that the aggregations are formed at the beginning of the dry 
season, persist through it and dissolve at the beginning of the wet season, 
plus the presence of excessive fat tissue and the absence of eggs in the 
females, seems to point to a case of aestivation in a state very close 
at least to diapause. Individual diapause would serve the purpose of 
living through the dry season by itself, but the communal aestivation 
would have the additional advantage of keeping the sexes together, thus 
guaranteeing an effective and early encounter, optimizing the chances 
of early copulations and consequently the production of fertile eggs when 
weather conditions are once again favorable for larval development. 

The fact that individuals abandon the groups from time to time, and 
the fact that we have witnessed individuals feeding at tree wounds in 
the neighboring woods, seem to indicate that this is not a case of com- 
plete diapause, but a partial one that calls for a close and reliable 
source of food, even if only periodically needed by organisms whose 
metabolism is grealty slowed down. 

There seems to be a degree of organization in the groups with some 
kind of discrimination between individuals belonging to different ones. 
The organization is suggested by the consistent way the groups are 
formed: there is a nucleus of several individuals with the heads pointing 
inwards, sometimes so close as to have their upraised antennae almost 
touching, surrounded by tightly packed rows, forming circles or partial 


Figs. 1 and 2. Roosting assemblies of Smyrna karwinskii: 1, 123 adults; 2, 39 
adults. Both photographed at Cerro Verde, 23 December 1972. 

circles of individuals with the heads again pointing inwards, and with 
the antennae touching a member of an inside row (see Fig. 1). The 
discrimination is deduced from the results of the disruption of the 
large group when it was counted: none of the dispersed individuals 
came to join the members of the smaller group, but kept by themselves. 
Probably they later formed another group elsewhere composed of the 

VOLUME 28, NUMBER 3 229 

same individuals. One month after the smaller group was partially 
disturbed, it again had about the same number it had had originally, 
including the one individual marked with an unfamiliar color. 

It is our opinion that Smyrna karwinskii adults have acquired this 
unusual social behavior as an adaptation tending to minimize the losses 
of individuals through the dry season resulting from predation on the 
one hand and excessive activity on the other, and to maximize the 
chances of early egg production when conditions are favorable for the 
dispersal of the species. This adaptive strategy nevertheless seems to 
be disadvantageous when compared with the one adopted by S. 
blomfildia datis, whose larval polymorphism seems to indicate a more 
flexible ability to adapt itself to adverse ambient conditions. 


We are greatly indebted to Dr. Alexander B. Klots of the American 
Museum of Natural History, who besides encouraging the authors to 
publish the results of their observations, took time out of his busy 
schedule to read the manuscript and give much valuable criticism. We 
also thank Dr. Allen M. Young who shared important information with 
us, Viktor Hellebuyck who helped the authors in parts of their observa- 
tions, and the rest of the family Muyshondt for their sustained efforts 
in the study of the Salvadorian butterflies. 


CarpPeNTER, G. D. H. 1931. Acraeinae butterflies congregating in a small area 
for the night’s rest. Proc. Roy. Entomol. Soc. London 6: 71. 

CiEeNncH, H. K. 1970. Communal roosting in Colias and Phoebis (Pieridae). J. Lepid. 
Soc. 24: 117-120. 

Crane, J. 1955. Imaginal behavior of a Trinidad butterfly, Heliconius erato hydara 
Hewitson, with special reference to the social use of color. Zoologica 40: 167— 

Hesse, J. H. 1966. Fluid recycling in Agathymus aryxna (Megathymidae). J. 
Lepid. Soc. 20: 242. 

Jones, F. M. 1930. The sleeping Heliconians of Florida. Nat. Hist. 30: 635-644. 

McFar.aAnp, N. 1971. A specialized case of commural roosting in Pieris rapae 
(Pieridae). J. Lepid. Soc. 25:144-145. 

McNeEiL, F. A. 1937. Notes on the gregarious resting habit of Danaeus melissa 
hamata W. S. Maclery, in the Whitsunday Islands off the East coast of 
Queensland. Proc. Roy. Entomol. Soc. London 12:108. 

Poutton, E. B. 1931. The gregarious sleeping habit of Heliconius charitoniua, L. 
Proc. Roy. Entomol. Soc. London 6: 71. 



Ciumrorp D. FERRIS” 
College of Engineering, University of Wyoming, Laramie, Wyoming 82070 

Erebia callias Edwards is found in Asia (Iran, Mongolia), Siberia, and 
the Rocky Mountains of the United States. It is closely allied to Erebia 
tyndarus (Esper) of the Old World, with which it was thought to be 
conspecific, until de Lesse (1955) demonstrated that the two species 
have different numbers of chromosomes. The diploid number for callias 
is 30 and for tyndarus, 20. 

This butterfly is not well known to collectors in the United States, 
probably became of its restricted habitat. It flies in the treeless Arctic- 
Alpine Zone above 10,000’. It is usually found in grassy areas, but I 
have also taken it on rocky outcroppings and flying about gravel patches. 
Several collectors have observed callias virtually swarming on Guanella 
Pass, Clear Creek Co., Colorado (observed by J. D. Eff in 1962, C. D. 
Ferris in 1967, and by O. Otto in 1972 as reported in the News of the 
Kepid) Soe. la) March) 1973, p23) 


Holland (1898) reported callias from Colorado and New Mexico. 
Warren (1936) listed the same areas in North America. Ehrlich & 
Ehrlich (1961) list Colorado and Wyoming. Callaghan & Tidwell (1971) 
give Utah records. At the present time, callias is known from four states. 

The county records are listed below and state localities are shown in 
ariex, Ji 

Colorado: Chafee, Clear Creek, Grand, Hinsdale, Lake, Larimer, 
Park, Summit (Brown et al., 1957; C. J. Durden, in litt., 1973). 

Montana: Carbon (collected by author). 

Utah: Summit, Uintah (Callaghan & Tidwell, 1971). 

Wyoming: Fremont, Park, Sublette (Ferris, 1971). 

A search of the major U.S. museum collections has failed to turn up 
any specimens from New Mexico. It is quite possible that callias occurs 
in the high mountains of the northern part of that state. Holland may 

1 Published with ane approval of the Director, Wyoming Agricultural Experiment Station, as 
Journal Article JA 6 

2 Research AS nts Allyn Museum of Entomology, Sarasota, Florida. Museum Associate, 
Los Angeles County Museum of Natural History, Los Angeles, California. 

VOLUME 28, NUMBER 3 231 

Fig. 1. Collection sites (black dots) for E. callias in North America. 

have had specimens which are now lost, or he may have projected the 
range into New Mexico based upon the distribution in Colorado. 

One would also expect to find callias in the Snowy Range Mountains 
of Wyoming (Albany and Carbon Cos.) as what appears to be suitable 
habitat exists. To date, the insect has not been collected in this area. 
This is a strange situation as callias is abundant in spots to the north 



Fig. 2. “Normal” forms of E. callias: (a) ¢ Palmer Lake, 10,800’, Sublette 
Co., Wyoming, 1 August 1972, genitalia Fig. 4 (b); (b), (ce) 66, and (d) 
2, Guanella Pass, 11,665’, Clear Creek Co., Colorado, 11 August 1968. 

and south of this area. Other alpine species normally associated with 
callias habitat, such as Colias meadii Edwards and Parnassius phoebus 
ssp., are found in the Snowy Range. 

It seems strange that callias has not been reported from Canada or 
Alaska, since it occurs in Siberia. Perhaps it will turn up as more regions 
open to travel. It flies late in the season (early August) as alpine species 
go, when many of the other high altitude species have either ceased 
flying or are on the wane, and for this reason, may have been overlooked 
in some areas. 


Fig. 2 illustrates three normal males and 1 female of E. callias. The 
two FW ocelli are fully developed and there is a HW submarginal row 
of three ocelli. Normally the FW ocelli are well-pupiled, while the 
HW ocelli vary in pupil size. This is the usual form found in Colorado, 
Utah, and central Wyoming. 

Fig. 3 illustrates the variation in callias that occurs along the Montana- 

VoLUME 28, NUMBER 3 230 

C 7 

Fig. 3. E. callias from Beartooth Pass area, U.S. Hwy. 212, Carbon Co., Montana, 

45 August 1972: (a.,b.d,e) 64; (ce), (f) 22. Some 2 lack ocelli entirely. 
Genitalia of (e) in Fig. 4 (a). 

Wyoming border on the Beartooth Plateau. “Normal” forms are found, 
but the majority of the specimens collected from this region are atypical 

when compared with Colorado material (Type Locality: Mosquito Pass, 
Park Co., Colorado). 


el] fi 

Fig. 4. Genitalia: (a) ¢ shown in Fig. 3 (e); (b) 4 shown amelie tae 
(ce) $ from Beartooth Pass area, Carbon Co., Montana, 5 August 1972; (d)-(f), 
Guanella Pass, Clear Creek Co., Colorado, 11 August 1968. 

With respect to the dorsal wing surfaces, all ocelli are absent in 
the extreme case, and only a fulvous patch appears on the FW. In 
most of the specimens, the HW ocelli are lacking. Only the pupils 
occur in other examples (FW) and the surrounding dark iris is absent. 
Other specimens exhibit FW ocelli that are substantially diminished in 

VoLUME 28, NUMBER 3 235 

size, and in some cases, the pupils are reduced to the point of 
obsolesence. Although subspecific names have been applied to E. 
callias in the Old World, it does not appear reasonable to propose 
another taxon for the Beartooth Plateau segregate. E. callias is a 
highly variable insect in both facies (Figs. 2, 3), and in genitalia 
(Fig. 4). There is no firm character, other than geography, upon 
which to erect a new taxon. B. C. S. Warren (pers. comm.) concurs 
in this matter. Warren (1936, p. 303) has also commented on the 
genitalic variation in the Old World races, and has identified two 
clasper types. 

No explanation is offered regarding the variation in the U.S. popu- 
lations. A parallel situation occurs with E. tyndarus in Europe and 
the non-ocellated form was described by Westwood (1851) as an 
aberrant vesagus. The vesagus form of tyndarus occurs locally as 
a form and in “normal” populations of tyndarus infrequently as an 
aberrant. In facies, tyndarus f. vesagus is identical with the Beartooth 
Plateau non-ocellated E. callias.. Warren (1936) figures French ma- 
terial of vesagus (Plate 89). Perhaps the form name vesagus could be 
applied to Wyoming-Montana callias, but infrasubspecific names have 
no standing in the I. C. Z. N. Code. 


The author would like to thank the following for supplying in- 
formation about E. callias in North America: Julian P. Donahue, Los 
Angeles County Museum of Natural History, Los Angeles, California; 
C. J. Durden, Texas Memorial Museum, Austin, Texas; Patrick J. 
Conway, Chicago, Illinois (for checking the Chicago Field Museum 
collection); Harry K. Clench, Carnegie Museum, Pittsburgh, Pennsy]l- 
vania; Dr. Frederick H. Rindge, American Museum of Natural History, 
New York, New York; Dr. Edwin M. Perkins, Jr., University of Southern 
California, Los Angeles, California; Mike Toliver, Albuquerque, New 
Mexico; B. C. S. Warren, Folkestone, England. 


Brown, F. M., J. D. Err & B. Rorcer. 1957. Colorado Butterflies. Denver 
Museum, Denver. 

CALLAGHAN, C. J. & K. B. Tiowetx. 1971. A checklist of Utah butterflies and 
skippers. J. Res. Lepid. 10: 191-202. 

DE LessE, H. 1955. Nouvelles formules chromosomiques dans le group d Erebia 
tyndarus Esp. (Lépidoptéres, Satyrinae). Comptes rendues hebdomadaires 
des séances de l’Académie des Sciences 240: 347-349. 

EnriuicH, P. H. & A. H. Enriicu. 1961. How to Know the Butterflies. Brown, 
Dubuque, Iowa. 


Ferris, C. D. 1971. An annotated checklist of the Rhopalocera of Wyoming. 
Sci. Monogr. 23. Agri. Exp. Stat., Univ. of Wyoming, Laramie. 

Ho.LiLanp, W. J. 1898. The Butterfly Book. Doubleday, New York. 

WarREN, B. C. S. 1936. Monograph of the Genus Erebia. British Museum 
(Natural History), London. 

Westwoop, J. O. 1851. In Doubleday, Gen. diurn. Lepid. 2: 380. 


Re-reading a note by Pyle (1972, J. Lepid. Soc. 26: 261) on a Lorquin’s 
Admiral (Limenitis lorquini burrisonii Maynard) that chased after a Glaucous- 
winged Gull brought to mind some observations I made this past summer in 
Bartlesville, Washington County, Oklahoma on bird and insect-chasing by Polygonia 
interrogationis (Fabricius). 

On 8 July 1973 I was exploring a field that contained a few trees surrounding 
a small marsh. Chimney Swifts (Chaetura pelagica) were often observed hunting 
over this area, sometimes making passes within 6 ft. of the ground surface. When 
a swift passed near a particular tree I noticed a butterfly in pursuit for some 20-— 
30 ft. before breaking off with the chase. The butterfly, a male Question Mark, then 
returned to the tree and began a methodical patrol of one section of the tree, flying 
back and forth in front of it with periodic darts and chases after other flying insects 
including beetles, dragonflies, and other butterflies. I sat down next to the tree and 
decided to observe the butterfly’s behavior, when another swift flew by. The 
Question Mark immediately took pursuit as before. After a brief chase the butterfly 
returned to the tree and resumed its patrolling. I was able to observe this behavior 
for several days but only in the evenings after 1800 hrs. At this time the butterflies 
were out patrolling and the Chimney Swifts were hunting over the field and marsh. 

Pyle suggested that the chasing behavior exhibited by his Lorquin’s Admiral was 
most likely a courtship chase, presumably the pursuit of a possible female. This 
possibly applies to the Question Mark since any flying object was pursued until 
it was apparently recognized. Another possibility is that these animals are exhibiting 
aggressive territorial behavior and are attacking all flying intruders. I observed some 
prolonged chases by two of the Question Marks in which they flew head-on at each 
other and beat the opponents wings with their own. Usually, however, one would 
make a “sneak attack” on the other and pursue it from the rear until it either chased 
the first temporarily away or was out-maneuvered. A third possibility would be a 
combination of the first two in which the butterfly leaves its post in pursuit of 
a possible female. When the butterfly identifies the object it either ceases pursuit 
or continues after in either an aggressive attack or a courtship chase. 

D. Paut HEenpricks, 305 East Maplewood Avenue, Littleton, Colorado 80121. 

VoLUME 28, NUMBER 3 Levy 


JouHn H. Masters 
5211 Southern Avenue, South Gate, California 90280 

Biennialism in insects is that situation where the insects life-cycle 
takes two years to complete and imagos are produced but once every two 
years. It may be accompanied by biennial-flights, when in a given locality 
adults fly only in alternate years, or it may be accompanied by annual- 
flights. Unless biennial-flights are involved, biennialism is very difficult 
to perceive in nature without carefully working out the life-histories. 
Annual-flights may occur when the species is only partially biennial or 
when two allochronic populations are involved. 

Many species of butterflies occurring in desert or near-desert regions 
are partially biennial. Papilio rudkini Comstock (Papilionidae), for 
example, is normally annual, but in especially dry seasons a portion of 
the population will remain in the pupal stage for an extra year before 
emerging. This is undoubtedly an advantage to the species as it reduces 
the risk of having an entire population wiped out in a particularly bad 
drought year. 

Recognized cases of regular biennialism, however, are very rare 
in Lepidoptera and they are confined to species (almost exclusively 
Satyridae) that occur in arctic, alpine or at least boreal regions; sug- 
gesting that biennialism may be their means of coping with very short 
growing seasons. In the Palearctic Region, species with proven bien- 
nialism are restricted to Oeneis jutta (Hubner) and several species of 
Erebia, including Erebia claudina (Borkhausen) and Erebia ligea (Lin- 
naeus ); although a large number of Erebia and several other Oeneis are 
suspect. In the Nearctic Region, five species of the genus Oeneis ( jutta, 
macounii (Edwards), nevadensis (Felder & Felder), chryxus (Double- 
day) and taygete Geyer) are known to be biennial in at least part of 
their ranges; several other species, including Erebia disa (Thunberg) 
(Masters, 1969), Erebia theano (Tauscher) (Masters, 1971) and Boloria 
polaris (Boisduval) (Nymphalidae) (Masters, 1971), are highly suspect. 

The best known example of polyennialism in insects is the “Periodical 
Cicada” or “Seventeen-Year Locust,” Magicicada septendecim (Linnaeus ) 
(Homoptera), which has a seventeen-year life-cycle that produces adult 
insects Once every seventeen years. Quite a few “broods” of M. 
septendecim are recognized, however, with each brood occupying a 
restricted geographical area distinctly different from other broods, and 


with each brood making its emergences as imagos on its own seventeen- 
year cycle. 

Biennialism does not produce a picture nearly as complex as septen- 
decennialism. However, in the genus Oeneis biennialism is usually ac- 
companied by geographic brood territories. In most cases populations 
over extensive areas are on the same brood-cycle and alternation with 
another completely allopatric population on the alternate brood-cycle 
occurs only across a natural barrier such as a mountain range or desert. 
In Oeneis (e.g. Oeneis jutta) the areas of biennial alternation frequently 
correspond to areas inhabited by different subspecies; these subspecies 
are both allopatric and allochronic. When two biennial species of Oeneis 
inhabit the same region, although not necessarily the same habitat, they 
invariably alternate with each other and display very pronounced bien- 
nialism. In the genus Erebia, biennialism results in biennial flights in 
which nearby colonies randomly alternate with each other on the year 
of flight. 

The types of O. macounii were collected at Nipigon, Ontario by 
Professor John Macoun in June 1884. The new species created quite a 
bit of interest and a number of persons journeyed to Nipigon to collect 
it, but with very mixed success. James Fletcher sought it in 1886, but 
got there in August and was too late for it, and again in 1887 which 
is the off year at Nipigon. Fletcher returned to Nipigon in 1888 with 
Samuel Scudder and was finally successful in getting it on July 5th. 
Fletcher wrote (1888) “I had been to Nepigon [sic! Nipigon] once before 
at exactly the right season and again a month later, but had not seen 
a specimen, and had begun to think that perhaps after all there might 
possibly be some mistake about the locality.” Oeneis macounii was not 
taken for several more years at Nipigon, but Alberta specimens turned 
up and interest gradually diminished in the Nipigon colony. 

As late as 1942, biennialism by Oeneis macounii was still not ‘suspect. 
George Shirley Brooks had a summer cabin at Victoria Beach, Manitoba 
where he for years collected large numbers of O. macounii for exchange or 
sale. He wrote (1942) “Oeneis macouni [sic! macounii] Edw. has been 
taken only in a limited area at Victoria Beach where it flies at irregular 
periods among Pinus banksiana. One year it may be abundant, and 
then it may be abundant, and then it may not be seen for several years.” 
Since all of Brooks specimens were taken in even-numbered years, it 
is surprising that he did not tumble onto the biennialism in this species— 
he collected it over a twenty year period at Victoria Beach. 

The fact that Oeneis macounii is biennial was well known by the 1960's, 



_ SOLOPLLI} POOIG,, PUB SJUSTF [VIUUSIG BSUNVSNIE ‘sisuappaau slauaQ pue NunodDUL Slaua_G Fo seBuRY “T “SI 


*“saeok pezoqunu-usAs ut 3UuTATF ‘TT poorq 

‘saeoA pezequnu-uere ut 8uTATF SII poorzq STINNOOVW STHNG0 C) it 

po 4h y) 




@ flying in odd-numbered years. 

© flying in even-numbered years. 

a flying in equal numbers every year. 

© flying every year, but in greater 
numbers in odd-numbered years. 

© Ne 
C) flying every year, but in greater 
numbers in even-numbered years. 
O ~ 
- \ 
4 ‘s 
= SCONSIN a an or aX 
kK. © 1 WI Rese a A 
: A 1 
A ; 
s A iB = 
x ; ; 
Ga @ 

Fig. 2. Distribution of the genus Oeneis in Minnesota, Wisconsin and Michigan, 
illustrating alternation of annual flights. 

but the first published record was by Masters, Sorenson & Conway (1967). 
C. S. Quelch first pointed out to me in 1966 that O. macounii colonies in 
Eastern Manitoba were on even-year cycles while those in Western 
Manitoba were on odd-year cycles. Since that time I have been gathering 
distributional and chronological data for all Oeneis species in order to 
demonstrate the point. 

In my map (Fig. 1) the known localities for Oeneis macounii and 
the closely related Oeneis nevadensis (Felder & Felder) are shown. It is 
readily apparent that three distinct “brood territories” exist. The break 
between Oeneis nevadensis and O. macounii is the Rocky Mountains. 
The break between the eastern, even-year brood and the western odd- 
year brood of O. macounii is Lake Winnipeg and the Red River Valley, 
which is the former location of Glacial Lake Agassiz. The southernmost 
localities for Oeneis nevadensis are for subspecies iduna (Edwards) 
which apparently flies annually. The allochronic eastern and western 
populations of Oeneis macounii have been isolated from each other at 
least 18,000 years, since before Lake Agassiz was formed in the late 
Pleistocene, however they exhibit no phenotypic distinctions that would 
warrant the designation of subspecies. 

VoLUME 28, NuMBER 3 241 

Oeneis jutta occurs through most of the range occupied by Oeneis 
macounii and is also a biennial species. The most remarkable circum- 
stance about this is the fact that O. jutta has the most pronounced 
biennialism where it comes into the same range as O. macounii and it 
alternates years with Oeneis macounii. This can be seen in the map 
(Fig. 2) showing a portion of the area where the two species are 
sympatric. The same dividing line (Lake Winnipeg—Red River Valley) 
that divides the eastern and western populations of Oeneis macounii, 
separates an eastern odd-year cycled population (subspecies ascerta 
Masters & Sorensen) from a western even-year cycled population (sub- 
species ridingiana Chermock & Chermock). In Minnesota where both 
species occur together, O. jutta is religiously biennial and can be taken 
only in odd-numbered years. In Wisconsin, east of the range of O. 
macounti, O. jutta may be taken in any year but exhibits a very strong 
population “pulse” occurring in vastly greater numbers in odd-numbered 
years. The further east you go, which is more distant from the range 
of O. macounii, the weaker this. pulse becomes. 

It would be attractive to theorize that interspecific competition has 
created the alternation in the annual flights of these two species, but 
this does not seem likely. For one thing, the two species have com- 
pletely different habitats; O. jutta occupies sphagnum-moss/black spruce 
bogs while O. macounii inhabits sandy ridges where jack pine grows. 
Both species are territorial and have very similar adult behavior, how- 
ever. A thorough discussion of the bionomics of these two species is 
given by Masters & Sorenson (1969). 

Other species of Oeneis that occur in the Lake Superior region include 
Oeneis uhleri varuna (Edwards) and Oeneis chryxus strigulosa Mc- 
Dunnough (Fig. 2). O. uhleri is a prairie inhabitant which apparently 
occurs every year in fairly equal numbers. O. chryxus strigulosa occurs 
southeast of the range of O. macounii, apparently having a habitat as- 
sociation with sedimentary rocks. O. chryxus flies every year in Michi- 
gan, but exhibits a strong “pulse” with much more pronounced flights 
in even-numbered years—thus alternating with O. jutta. 


Brooks, G. S. 1942. A check list of the butterflies of Manitoba. Can. Entomol. 
7A: 31-36. 

FLETCHER, J. 1888. A trip to Nepigon, some notes upon collecting and breeding 
butterflies from the egg. 19th Annual Report, Entomol. Soc. Ontario: 74-88. 

Masters, J. H. 1969. Ecological and distributional notes on Erebia disa (Satyridae ) 
in central Canada. J. Res. Lepid. 7: 19-22 [1968]. 


. 1971. Butterflies of Churchill, Manitoba. Mid-Continent Lepid. Series 
25: 1-16. 

& J. T. SorENsEN. 1969. Field observations on forest Oeneis (Satyridae ). 
J. Lepid. Soc. 23: 155-161. 

, J. T. SorENSEN & P. J. Conway. 1967. Observations on Oeneis macounii 
in Manitoba and Minnesota. J. Lepid. Soc. 21: 258-260. 


A collecting trip to the Providence Mountains of eastern San Bernardino County, 
California, on 18 May 1973, resulted in a surprise capture—a new state record. 
While hiking down the south fork of Bonanza King Mine Canyon, I spotted two little 
dark butterflies flying close to the streambed. Both were captured at 1530. Con- 
firming my initial suspicions, they were two males of Phyciodes texana (Edwards). 
One was in fairly good condition, the other rather worn. The two captured speci- 
mens were the only texana sighted that afternoon. This appears to be the first 
recorded capture of this species in California. It was not figured in any of the 
older books on California butterflies. And it was not mentioned in the recently 
published book, The Butterflies of Southern California by Thomas C. Emmel and 
John F. Emmel (Los Angeles County Museum of Natural History and the Ward 
Ritchie Press). 

Captures of Arizonan butterflies are not unusual in the mountains of eastern 
San Bernardino County. Although part of the Mojave Desert, the Providence 
Mountains, the Ivanpah Mountains, the New York Mountains and the Sacramento 
Mountains tend to resemble in fauna and flora the Sonoran Desert of southern 
Arizona. These ranges adjacent to the Colorado River Valley receive more rainfall 
than the lower portions of the Mojave Desert. And they have a rainfall distribution 
similar to the Sonoran Desert, with winter rains being supplemented by thunderstorms 
in the summer. It is not surprising, therefore, that we get occasional reports of Phoebis 
sennae (Linnaeus), Phoebis agarithe (Boisduval), Colias caesonia (Stoll), Eurema 
mexicana (Boisduval), Limenitis bredowii eulalia (Doubleday) and Strymon colu- 
mella (Fabricius) from this region. Lepidopterists should be on the lookout for 
other Arizona butterflies straying into this area. This unique region may very well 
produce other new state records. 

RicHARD C. PriesraF, 5631 Cielo Avenue, Goleta, California 93017. 

VoLUME 28, NUMBER 3 243 


Route 4, Box 104-EB, San Antonio, Texas 78228 

Two species of moths representing two families are recorded here 
as new to the United States. Only reared examples are known for one 
of these species, the other is represented by a single field collected 
example. Neither species is known to be of any economic importance. 

Gnophaela aequinoctialis (Walker ) 

Dioptis aequinoctialis Walker, 1854. List of Specimens of Lepidopterous insects 
in the Collection of the British Museum. Lepidoptera Heterocera. London. 2: 
331. (TL: South America). 

While collecting in Panther Canyon above Landa Park, New Braunfels, Comal 
County, Texas, W. W. McGuire took one female on 9 April 1972. This specimen 

] ? 

Fig. 1. Gnophaela aequinoctialis (Walker), 2, dorsal view; New Braunfels, Texas. 
(W. W. McGuire). Wing expanse 52 mm (center of thorax to tip of FW x 2). 

1 Contribution No. 282. Bureau of Entomology, Division of Plant Industry, Florida Department 
of Agriculture and Consumer Services, Gainesville 32602. 

2Research Associate, Florida State Collection of Arthropods, Division of Plant Industry, 
Florida Department of Agriculture and Consumer Services. 


4 4 

Figs. 2 & 3. Elymiotis notodontoides Walker, ¢, @ respectively, dorsal view; 
McAllen, Texas. (Roy O. Kendall). Wing expanse, ¢ 40 mm, 2 50 mm (center 
of thorax to tip of FW x 2). 

is now in the collection of André Blanchard of Houston, Texas who kindly 
photographed it for illustration (Fig. 1). 

There are twenty-six examples of this species represented in the National Mu- 
seum of Natural History, Washington, D.C., all from Mexico. Specific data 

VoLUME 28, NuMBER 3 245 

are: twenty-one, Popocatepetl Park [Puebla], Mexico, 8,000’, June; one, Jalapa 
[Oaxaca], Mexico; one, Tehuacan [Puebla], Mexico; two, Mexico; one, Cuernavaca 
[Morelos], Mexico. No other data available. The nearest of these locations is about 
900 air miles from where the U.S. specimen was taken. 

Elymiotis notodontoides Walker 

Walker, F., 1857. List of Specimens of Lepidopterous Insects in the Collection 
of the British Museum. Lepidoptera Heterocera. London. 11: 609. Described 
from a single male from an unknown locality. 

On 11 September 1972, the author collected 4 last instar larvae feeding on the 
foliage of Malpighia glabra L., Malpighiaceae, a native plant of the area, but in 
this instance growing as an ornamental at a motel in McAllen, Hidalgo County, 
Texas. Further examination disclosed 9 pupae and 8 empty pupal cases at the 
base of this small shrub. Pupation took place in loose cocoons constructed in 
dried leaves and debris. Two larvae, two pupae, and all empty pupal cases were 
preserved in alcohol. The two remaining larvae pupated 14 and 17 September; 
adults, both males, emerged 24 and 27 September. Other adults emerged, ex 
pupae: 18-IX-72 (9), 19-IX (4), 20-IX (16, 19), 21-IX (14, 19), and 22-Ix 
(6), for a total of 64, 39 including the two ex larva. A male and female are 
illustrated (Figs. 2 & 3). 

The pair which emerged 20 September were found in copula about 0730. The 
female was kept alive for egg production. Eighteen eggs were found in the con- 
tainer 22 September, only two of which were deposited on the larval foodplant. 
All of these eggs were preserved. The female was fed and placed over a caged 
living larval foodplant in the laboratory garden. After returning from a field 
trip on 30 September, the female was not to be found. No doubt it had died 
and had been eaten by ants. Only two eggs could be found, and these on the 
foliage. Near the end of October, examination disclosed the eggs had not hatched. 
Assuming the eggs to be fertile, and pending further study, it would appear this 
species may have an ovum diapause. 

Examples of this species from other locations in the National Museum of Natural 
History are: three, Paso San Juan, Veracruz, Mexico; one, Coatepec [Veracruz], 
Mexico; one, Tepic [Nayarit], Mexico, June; one, Cabezas nr. Cardel, Veracruz, 
Mexico, July; one, Cajuli Sbo Domingo; one ¢, Constanza, Dominican Republic, 
2-6 June 1969, leg. Flint & Gomez; one 2, Los Hidalgos, Dominican Republic, 4— 
5 June, leg. Flint & Gomez. This species has not been collected on the other islands 
of the Antilles by any of the N.M.N.H. recent collectors. 


I wish to thank Dr. E. L. Todd, Systematic Laboratory, U.S. Depart- 
ment of Agriculture, at the National Museum of Natural History, for 
determining these species, for providing data on like specimens in the 
national collection, and reference citations. I’m also indebted to André 
Blanchard for photographing the specimens illustrated. 



Epwarp C. KNUDSON 

Department of Radiology, Baylor College of Medicine, 
Texas Medical Center, Houston, Texas 77025 

One hears so much about the population explosion in Florida, that 
it seems difficult to imagine at times how even a butterfly could survive 
it. Indeed, the drainage of the swamps, burning and bulldozing of 
forest lands, and increasing levels of pollution have gravely endangered 
many of Florida’s unique species of Lepidoptera. In spite of this, 
certain other species are experiencing a ‘population explosion’ of their 

One such species is Urbanus dorantes dorantes Stoll, which, prior to 
1969, was not believed to be present in Florida. Kimball (1965) listed 
an old record from the Cleveland Museum, labeled Tampa, 1908, and 
Miami, 1916, but this was thought to be fraudulent. However, Clench 
(1970) discovered that the species was common at Chokoloskee (Collier 
County) on 20 November 1969. During March and April 1970, Miller 
& Miller (1970) took specimens at Homestead (Dade County), Key 
Largo, and Tavernier (Monroe County). Pliske (1971) reported that 
U. dorantes dorantes was common in Dade County from November 
1969 through January 1970. Also, C. Hallas has reported specimens 
from Dade County in April and December 1971, and from Key Largo in 
January 1972. In addition, Burris (1973) took specimens in Hillsborough 
County during February and April 1972. 

The first specimen of U. dorantes dorantes taken by the author was 
found in Pahokee (Palm Beach County) on 12 June 1971. Then, on 
12 October 1972, dorantes was found to be common near Bartow (Polk 
County). A more careful search of more northerly areas revealed this 
hesperiid to be present in Gainesville (Alachua County) as well. 
Urbanus proteus L. was abundant at the time and it was estimated 
that U. dorantes dorantes comprised at least five percent of the tailed 
skippers seen in the Gainesville area. On 16 October 1972 several 
specimens were taken in Yulee (Nassau County) at blossoms of iron- 
weed and Trilisa sp. along the roadside. 

On 4 November 1972 U. dorantes dorantes was found to be common 
at Lake Worth (Palm Beach County), Homestead, and Key Largo. 
In these localities it was clearly the dominant tailed skipper. On Key 

VoLUME 28, NUMBER 3 247 

Largo a half acre field, overgrown with Lantana sp. and Bidens pilosa 
L., was estimated to contain about one hundred specimens of dorantes 
and only a few of proteus. The flight pattern of the two seemed to be 
quite similar, although dorantes was somewhat faster and less erratic 
than proteus. The absence of green iridescence in dorantes was not 
easily evident while on the wing, and the two species were best distin- 
guished in the field by the difference in maculation of the underside 
of the hindwing. On upper Key Largo, U. dorantes dorantes could also 
be found along the roadside and on shaded trails through the tropical 
hardwood forest. Along the forest trails it was observed that the flight 
pattern of dorantes was remarkably similar to that of Polygonus leo 
(Gmelin), i.e., dorantes would dart back and forth between the dense 
vegetation on either side of the trail and finally alight on the under- 
side of a leaf. 

U. dorantes dorantes was again observed in the same localities in 
southern Florida during late November through December 1972, and 
on 18 December 1972 it was captured in Largo and Dunedin (Pinellas 
County). Subsequently the populations declined, although on 6 March 
1973 dorantes was still present on Key Largo and was also found at 
Devils Gardens (Hendry County ). 

During October, November, and December 1972, a careful search 
was made for the larvae of U. dorantes dorantes on leguminous plants 
at various locations. These plants were: at Bartow, Pueraria thun- 
bergiana (S. & Z.); at Lake Worth, Vigna marina Merrill, at Homestead, 
Glycine max L. and Phaseolus lathyroides L.; and at Key Largo, Galactia 
spiciformis Torr & Gray and Desmodium tortuosum DC. The plants 
were checked by hand and by the use of a D-Vac (back-pack suction 
machine). However no larvae of dorantes were found. 

In summation, it appears that for the past three years U. dorantes 
dorantes has been common in southern Florida, with a peak abundance 
during November, December, and January. It also appears that in 1972 
dorantes extended its range far northward along the coast and into 
the interior sections. All specimens taken in Florida belong to the 
subspecies dorantes (distinguishing characteristics may be found in 
Clench (1970) ). This fact implies that dorantes was not an introduction 
from Cuba, as a distinct subspecies occurs there. However, U. dorantes 
dorantes is common in southeastern Texas ranging as far north as the 
Dallas area, and thus conceivably it could have reached Florida from 
a northern route, around the Gulf coast. If so, the records from 1908 
and 1916 are perhaps valid after all. It seems incomprehensible, how- 
ever, that dorantes could have been overlooked for fifty years, especially 


in view of the extensive collecting in south Florida. One is also at a 
loss to explain the absence of dorantes along the northern Gulf coast. 

The most logical explanation, Miller & Miller (1970), is that dorantes 
was introduced artificially into southern Florida, or possibly transported 
to the area by the winds of Hurricane Camille in August 1969. Because 
of its range in Texas, dorantes may well prove to be a late summer 
visitor throughout northern Florida and coastal Georgia. 


I wish to thank T. M. Neal for his assistance and many valuable 
observations. Also, I am grateful to L. D. Miller, H. V. Weems, C. P. 
Kimball, and C. Hallas for their observations, comments, and suggestions 
that led to this paper. 


Burris, D. L. 1973. Interesting Florida butterfly records. J. Lepid. Soc. 27: 84. 

CiLencu, H. K. 1970. New or unusual butterfly records from Florida. J. Lepid. Soc. 
24 QAQ=244, 

KIMBALL, C. P. 1965. Lepidoptera of Florida. Div. Plant Industry, Gainesville. 

Kuots, A. B. 1951. A Field Guide to the Butterflies. Houghton Mifflin, Boston. 

Miuter, L. D. & J. Y. Mitxier. 1970. Pieris protodice and Urbanus dorantes in 
southern Florida. J. Lepid. Soc. 29: 244-247. 

Putske, T. E. 1971. Notes on unusual species of Lepidoptera from southern Florida. 
J. Lepid. Soc. 25: 294. 


During the summer of 1973 I came across two instances of Speyeria spp. being 
attracted to the amber-colored glass used on the signal lights of motor vehicles. 

The first instance involved my motorcycle. I had left the machine parked on 
a disused logging road on Mt. Sicker, Vancouver Island. I returned just in time to 
see a butterfly alight on one of the amber lights. On approaching I saw that 
it was quivering its wings rapidly in the manner often seen when a male butterfly 
has settled near a receptive female. Its attention was completely focused on the 
colored glass, and I netted it easily. It was a male Speyeria hydaspe Bdv. 

In the second case a pickup truck was the attraction. It was parked near the 
summit of Mt. Prevost, the butterfly circled it several times, on each circuit dipping 
towards each of the little amber lights. This insect was quite wary, and I failed 
to collect it. It was a Speyeria, either S. hydaspe or S. zerene, as these are the 
only two species occurring in the vicinity. 

RicHarp Guppy, Thetis Island, British Columbia, Canada. 

VoLUME 28, NUMBER 3 249 

JNINID) Wield, WINIMaID) ube Mas: 

Route 4, Box 104-EB, San Antonio, Texas 78228 

The object of this paper is to remove the dubious status of earlier 
reports of two species of Lepidoptera being found in Texas. Each 
species is represented at present by a single example only. Examples 
of earlier recordings have not been found; it is possible, however, that 
they do exist. 

These species may represent single-brooded migrants which come to 
Texas from time to time. A precise judgment on this cannot be made 
until life history studies are conducted. Such studies would disclose 
critical ecological influences upon each. Another possible conclusion is 
that they are actually established in our fauna, but at such low popu- 
lation levels that they are seldom encountered by collectors. In any 
event, based on the good condition of these particular examples, we may 
conclude that they had not been on the wing long. No major climatic 
disturbances were involved. 

Enantia melite (Linnaeus) 1763 

Papilio melite Linnaeus, 1763. Amoen. Acad., vol. 6, p. 403 (gives habitat as 
Indiis ). 

Leptalis melite Linnaeus, 1767. Syst. Nat., 775; Skinner, 1898, A Syn. Catalogue 
of N. A. Rhopalocera; Dyar, 1902, A list of N. A. Lepidoptera (gives distribution 
as Mexico, New Mexico). 

Dismorphia melite: McDunnough, 1938, Check List of Lepid. of Canada and the 
U.S.A. (lists as doubtful N. A. occurrence); Holland, 1955, The Butterfly Book 
(credits to our fauna on the authority of Reakirt). 

Licinia melite: Klots, 1951, A field Guide to the Butterflies (vaguely recorded from 
Texas ). 

Enantia melite: Ehrlich & Ehrlich, 1961, How to Know the Butterflies (may oc- 
casionally stray across our southern border); dos Passos, 1961, J. Lepid. Soc. 15: 
211 (of doubtful occurrence in the Nearctic region). 

One example of this species was collected 3 September 1972 by 
W. W. McGuire in Bentsen-Rio Grande Valley State Park, Hidalgo 

County, Texas. The specimen, illustrated in Fig. 1, is in McGuire’s 

1 Contribution No. 289. Bureau of Entomology, Division of Plant Industry, Florida Department 
of Agriculture and Consumer Services, Gainesville 32602. 

2Research Associate, Florida State Collection of Arthropods, Division of Plant Industry, 
Florida Department of ‘Agriculture and Consumer Services. 



Fig. 1. Enantia melite (Linnaeus), ¢, dorsal view; Hidalgo County, Texas. 
(W. W. McGuire). Wing expanse 54 mm (center thorax to tip FW xX 2). 

Hamadryas amphinome mexicana (Lucas) 1853 

Papilio amphinome Linnaeus, 1767. Syst. Nat. (X), i, p. 473, no. 95. 
Hamadryas Hubner, 1806. Samml. Exot. Schmett. 1 pl. [47]; selected Papilio 
amphinome L. as the type species. Hemming, 1934, The Generic Names of the 
Holarctic Butterflies, vol. 1 (1758-1863), British Museum (N.H.), London, states 
that Ageronia, though valid nomenclatorially, is not required, as chloé Stoll, selected 
by Scudder as the type, is congeneric with amphinome L.; also, that Peridromia, 
also valid nomenclatorially, is not required as arethusa Cramer, selected by Scudder 
as the type, is congeneric with amphinome L., the type of Hamadryas Hb. 
Peridromia mexicana Lucas, 1853. Revue et magasin de zoologie, p. 311-312 (TL: 
Mexico ). 

Ageronia amphinome mexicana (Lucas): Frustorfer, 1913, in Seitz, Gross-schmett. 
Ende 5: 543-544, gives distribution as Texas, Mexico, Guatemala, Honduras, Chiriqui. 
Hamadryas amphinome mexicana: Monroe, Rose S., Garry N. Ross, and Roger N. 
Williams, J. Lepid. Soc. 21: 195, collected 2 ¢ at El Jaral, Honduras, 9 & 13 
August 1962. 

Although Frustorfer included Texas in the distribution for this sub- 

species, the name has never appeared on any check-list of Rhopalocera 
for the United States. In an attempt to locate other possible existing 


Fig. 2. Hamadryas amphinome mexicana (Lucas), 6, dorsal (2A) and ventral 
(2B) views; Hidalgo County, Texas. (W. W. McGuire). Wing expanse 76 mm 
(center thorax to tip FW x 2). 

VoLuME 28, NUMBER 3 


examples, several museums were contacted. In letter dated 24 August 
1973, Dr. P. Viette, Museum National D’histoire Naturelle, Paris, in- 
formed the writer that the type series of Peridromia mexicana could not 
be found. A letter dated 11 October 1973 from Mr. P. R. Ackery of the 
British Museum (N.H.), London, advised that no examples of mexicana 
could be found in the collection from locations farther north than 
Mexico. Mr. William D. Field, National Museum of Natural History, 
Washington, D.C., advised in letter dated 23 February 1973 that he 
could find no examples in the national collection labeled mexicana. 

One example of this subspecies was collected 3 September 1972 by 
W. W. McGuire in Bentsen-Rio Grande Valley State Park, Hidalgo 
County, Texas. The specimen, illustrated in Fig. 2, is in McGuire's 


Special thanks are extended to Mr. André Blanchard and his wife 
May Elise for photographing the specimens illustrated. I also wish to 
thank Dr. P. Viette, Mr. P. R. Ackery, and Mr. William D. Field for 
checking the collections in their charge and for other valuable informa- 


Klots (1951, A Field Guide To The Butterflies, Boston) listed Pellicia costimacula 
Herrich-Schaffer as occurring casually in the United States. In 1964, dos Passos (A 
Synonymic List of the Neartic Rhopalocera) dropped P. costimacula from the United 
States list and substituted P. angra Evans, a superficially similar species that was 
undescribed at the time Klots was published. 

I have taken fifteen male Pellicia in the Rio Grande Valley of Texas in the past 
six years. The genitalia of all correspond exactly to Evans’ figure (1953, Catalog 
of the American Hesperiidae, London. Part III, pl. 33) of P. costimacula arina 
Evans. Therefore, this species should be returned to the United States list. I am 
advised by H. A. Freeman (pers. comm.) that there is good justification for con- 
sidering arina to be a valid species, especially on the basis of genitalia. 

Mixes A. Rickarp, 4628 Oakdale, Bellaire, Texas 77401. 



Department of Biology, University of Louisville, Louisville, Kentucky 40208 

Kentucky has been one of the more poorly studied states with regard 
to its insect fauna. Until recently very little had been published on the 
Lepidoptera of the “Bluegrass State,” except for the many descriptions 
of new Microlepidoptera by V. T. Chambers of Covington (near Cin- 
cinnati, Ohio). The most recent state list of butterflies was that of Wheat 
(1908) [1909], which was poorly annotated and which included only 
65 species. Before that, Hattie Warner published two nearly identical 
lists (1894, 1895), the first with 60 species and the second with 61. All 
three of these early lists were based on specimens in the collection of 
the Kentucky Agricultural Experiment Station, Lexington. 

While Kentucky records are mentioned in various broader works on 
North American Lepidoptera, the state seems to be a blank on the 
distribution maps of most faunal and taxonomic publications. A few 
sources of information include more local coverages: Cook (1948) wrote 
of spring collecting in his area, and published the first record of Erora 
laeta from Kentucky. Merritt (1948) published the most exhaustive and 
complete list to that time, treating the fauna of Jefferson County (which 
includes Louisville). His work includes 63 species from that county, 
plus 6 more from within 100 miles. Records from an unpublished Master's 
thesis by D. K. Weniger (1946) augmented his own as source material. 
Covell (1969) provided a pre-impoundment list of one weekend's col- 
lecting in lowlands of Trigg Co. which are now beneath the waters of 
Lake Barkley. 

Additional printed information is to be found in the mimeographed 
“Club Notes” of the now-defunct Moth and Butterfly Club, which 
existed from 1946 to 1955. H. O. Ladd of Elizabethtown and James 
Unseld of Gravel Switch included some reports of Kentucky butterflies 
they had observed and collected. Finally, one may consult reports in 
the Field Season Summary of the Lepidopterists’ Society for the years 
1948-1952 and 1959-1972 (except 1962). Major contributors to these 
summaries include Carl Cook, James R. Merritt, Burt L. Monroe, Jr., 
Ralph Beebe, J. B. Wood, and myself. 

Since arriving at the University of Louisville in 1964, one of my projects 

1 University of Louisville Contributions in Biology No. 165 (New Series). 


has been to prepare a thorough annotated checklist of all Kentucky 
Lepidoptera. So far, of the more than 1350 species recorded from the 
“Bluegrass State” in my card file, only 123 are butterflies. It is the 
purpose of this paper to make known the species I know to have been 
collected or observed in Kentucky, with hopes that those who know 
of additions to this list will help extend the number by submitting their 
records. The broader work will hopefully be ready for publication in 
the next year or so. 

Another reason for publishing a butterfly list at this time is to end 
the confusion caused in the past by my circulating ditto-reproduced lists, 
occasionally revised, for the purpose of informing my colleagues of 
the known Kentucky butterflies. Some workers are desirous of having 
a published, up-to-date list to aid them in their revisionary and faunal 
studies of specific butterfly groups. 

Although the contributions of recent collectors in Kentucky will be 
included in the larger study in preparation in connection with the new 
records for which they are responsible, I would like to thank the fol- 
lowing for their cooperation in making this list possible: William R. 
Black, Jr., Annette F. Braun, Carl Cook, Carl C. Cornett, Charles V. 
Covell III, Charles J. Dempwolf, James K. Ettman, Greg Florence, Loran 
D. Gibson, Robert V. Gregg, James Harrod, Richard Henderson, L. C. 
Koehn, H. O. Ladd, James R. Merritt, Burt L. Monroe, Jr., Siegfried 
Scholz, J. A. Shields, James Tuttle, and J. B. and Lark Eyone Wood: 

Nomenclature used here follows the 1964 dos Passos Synonymic List 
of the Nearctic Rhopalocera and its two later partial revisions, with a few 
other changes reflecting more recent articles changing nomenclature and 
taxonomic status (as with Lethe species, and the use of Cynthia). 
The only annotation used here is the asterisk placed before the names 
of species for which I have only one or two records, or for which there 
seem to be acceptable sight records. While data on any species of 
Kentucky butterflies will be most welcome, I would especially like to 
hear of captures of species either missing from the list, or indicated with 
the asterisk. 

Family Hesperiidae 

Ambliscirtes samoset (Scudder) *Poanes viator (Edwards ) 
Ambliscirtes aesculapius (Fabricius ) Atrytone delaware (Edwards) 
Ambliscirtes vialis (Edwards ) Atalopedes campestris (Boisduyal ) 

* Atrytonopsis hianna (Scudder ) Pompeius verna (Edwards ) 

Euphyes vestris metacomet ( Harris ) Wallengrenia otho egeremet (Scudder) 
Poanes hobomok (Harris ) Polites coras (Cramer ) 

Poanes zabulon (Boisduval and LeConte) Polites themistocles (Latreille ) 


Polites origenes (Fabricius ) 
*Hesperia metea Scudder 

Hesperia leonardus Harris 

Hylephila phyleus (Drury ) 
Thymelicus lineola (Ochsenheimer ) 
Ancyloxypha numitor (Fabricius ) 
Lerema accius (Smith) 

Nastra lherminier ( Latreille ) 

Pholisora catullus (Fabricius ) 

Pyrgus communis (Grote) 

Erynnis icelus (Scudder and Burgess ) 
Erynnis brizo (Boisduval and LeConte ) 
*Erynnis baptisiae (Forbes ) 

*Erynnis zarucco (Lucas ) 

*Erynnis funeralis (Scudder and Burgess ) 
Erynnis martialis (Scudder ) 

Erynnis horatius (Scudder and Burgess ) 
Erynnis juvenalis (Fabricius ) 
Staphylus mazans hayhurstii (Edwards ) 
Thorybes bathyllus (Smith) 

Thorybes pylades (Scudder) 

Thorybes confusis Bell 

Achalarus lyciades (Geyer) 

Autochton cellus (Boisduval and Le- 

Conte ) 

*Urbanus proteus (Linnaeus ) 
Epargyreus clarus (Cramer ) 

Family Papilionidae 

Battus philenor (Linnaeus ) 
*Battus polydamas (Linnaeus ) 
Papilio polyxenes asterias Stoll 
Papilio cresphontes Cramer 

Papilio glaucus Linnaeus 
Papilio troilus Linnaeus 
*Papilio palamedes Drury 
Graphium marcellus (Cramer) 

Family Pieridae 

Pieris protodice Boisduval and LeConte 
*Pieris virginiensis Edwards 

Pieris rapae (Linnaeus ) 

Colias eurytheme Boisduval 

Colias philodice Godart 

Colias cesonia (Stoll) 

Phoebis sennae eubule ( Linnaeus ) 
*Kricogonia lyside (Godart) 

Eurema lisa Boisduval and LeConte 
Eurema nicippe (Cramer ) 

Nathalis iole Boisduval 

Anthocaris midea Hubner 

Euchloe olympia (Edwards ) 

Family Riodinidae 

Calephelis borealis (Grote and Robinson ) 

Family Lycaenidae 

Harkenclenus titus mopsus (Hiibner ) 

Satyrium liparops strigosa (Harris ) 

Satyrium calanus falacer (Godart ) 
*Satyrium caryaevorus (McDunnough) 

Satyrium edwardsii (Saunders ) 

Satyrium acadia (Edwards ) 

Calycopis cecrops (Fabricius ) 
*Callophrys irus (Godart) 

Callophrys henrici (Grote and Robinson ) 

Callophrys augustinus croesioides Scudder 

Callophrys niphon (Hiibner) 

Callophrys gryneus ( Hiibner ) 

Atlides halesus (Cramer ) 

*Eurystrymon ontario (Edwards ) 
Panthiades m-album ( Boisduval and Le- 
Conte ) 
Strymon melinus Hubner 
*Erora laeta (Edwards ) 
Feniseca tarquinius (Fabricius ) 
Lycaena thoe (Guerin-Meneville ) 
Lycaena phlaeas americana Harris 
Everes comyntas (Godart) 
Glaucopsyche lygdamus (Doubleday ) 
Celastrina argiolus pseudargiolus (Bois- 
duval and LeConte ) 


Family Libytheidae 
Libytheana bachmanii (Kirtland ) 

Family Nymphalidae 

Anaea andria Scudder Polygonia interrogationis (Fabricius ) 
Asterocampa celtis (Boisduval and Le- Polygonia comma ( Harris ) 
Conte ) *Polygonia faunus (Edwards ) 
Asterocampa clyton ( Boisduval and Le- Polygonia progne (Cramer) 
Conte ) Chlosyne nycteis (Doubleday ) 
*TLimenitis arthemis arthemis (Drury ) *Chlosyne gorgone (Hibner) 
Limenitis arthemis astyanax (Fabricius) | *Chlosyne harrisii (Scudder ) 
Limenitis archippus (Cramer ) Phyciodes tharos (Drury ) 
*Anartia jatrophae( Johansson ) Euphydryas phaeton (Drury ) 
Vanessa atalanta (Linnaeus ) Boloria toddi (Holland ) 
Cynthia virginiensis ( Drury ) Speyeria idalia (Drury) 
Cynthia cardui (Linnaeus ) Speyeria diana (Cramer) 
Junonia coenia (Hubner ) Speyeria cybele (Fabricius ) 
*Nymphalis vau-album (Denis and Schif- *Speyeria aphrodite (Fabricius ) 
fermiiller ) Euptoieta claudia (Cramer) 
Nymphalis milberti (Godart) Agraulis vanillae (Linnaeus ) 

Nymphalis antiopa (Linnaeus ) 

Family Danaidae 

Danaus plexippus (Linnaeus ) 

Family Satyridae 

Lethe anthedon (Clark) Euptychia hermes sosybius (Fabricius ) 
*TLethe creola (Skinner ) Euptychia cymela (Cramer ) 
Lethe appalachia Chermock Cercyonis pegala ( Fabricius ) 

Euptychia gemma (Hubner ) 


Cook, C. 1948. Early spring collecting in Kentucky. Lepid. News 2: 22. 

CovELL, C. V., Jr. 1969. Some pre-flood butterfly records from the Land Between 
the Lakes. Ky. Nat. 23: 8-9, 2 figs. 

Merritt, J. R. 1948. List of the Butterflies of Jefferson County, Kentucky. Annals 
Ky: Nat. Hist. I: 27-32. 

Warner, H. H. 1894. Kentucky butterflies. Can. Entomol. 26: 289-291. 

1895. A synopsis of the diurnal Lepidoptera of Kentucky. Dept. Zool. & 
Entomol. of State College, Lexington, Ky. 42 p. 

Wueat, F. M. 1908 [1909]. The diurnal Lepidoptera of Kentucky with key for 
their identification. Bull. State Univ. Ky. 1: iv + 173 p. 


VoLUME 28, NUMBER 3 25 


Department of Biology, Lawrence University, Appleton, Wisconsin 54911 

This report is one of a series of descriptive papers on the biology 
of species of ithomiid butterflies sympatric in one mountainous region 
of Costa Rica. It deals with the biology of Pteronymia notilla Butler & 
Druce. While a substantial amount of information is available on the 
taxonomy and phylogeny of the Central American Ithomiidae (Fox, 
1968), my reports (Young, 1972a,b) emphasize: (1) description of im- 
mature stages, (2) larval foodplant records, and (3) selected behavior 
patterns of larvae and adults. The eventual goal of these seemingly 
unrelated studies of different genera and species in the Ithomiidae is 
to describe the ecological and behavioral mechanisms that account for 
the observed local species diversity and structure of the ithomiid com- 
munity at the single locality selected for study. While we are a long 
way from this synthesis, the present paper includes the kinds of informa- 
tion on life history that will provide the foundation for interpretation of 
community structure. Inherent in this approach to the community ecology 
of tropical butterflies is the conviction that local community structure 
in these organisms is determined to a large extent by generic and species 
differences in adult reproductive behavior and larval foodplant selectivity. 


The selected locality is known as “Cuesta Angel” and is located on 
the Caribbean side of the Central Cordillera that runs through Costa 
Rica. The locality is about 8 km from Cariblanco (Heredia Province) 
and the specific area under study is one slope of a 300-meter deep forest- 
covered ravine (Fig. 1) whose bottom is the Rio Sarapiqui. The ridge- 
top elevation of the ravine is about 1000 m above sea level with per- 
sistent cloud cover (Fig. 1) and the general region is montane tropical 
forest or cloud forest. The ithomiine fauna is rich in both the forest 
understory and second-growth patches that are most abundant along 
a roadcut (the road to Puerto Viejo), which runs about 10 m from the 
ridge-top of the slope selected for study. Several genera of ithomiines, 
including Dircenna, Godyris, Oleria, Hymenitis, and Pseudoscada (in 
addition to Pteronymia) can be found both in river-bottom forest as 
well as ridge-top forest and second-growth. The river-bottom forest 


Fig. 1. The ravine at Cuesta Angel along the Central Cordillera in Costa Rica, 
where field studies of Pteronymia notilla were conducted. The butterfly is abundant 
in the understory of the forest down the sides of the ravine, and also along the 
edges of cleared second-growth (foreground and opposite ridge-top) associated with 
a readcut. 

is easily accessible via a small rock road that winds down the slope 
from the roadcut, and eventually goes up the opposite side of the ravine 
to the second ridge-top, where a small farming colony is located (see 
the cleared area on top of the opposite side of the ravine in Fig. 1). 
Other areas along the slope of the ravine are accessible with the use 
of rope and harness to work down the mountain side between roadcuts. 

Most of the field observations on adults and immatures of P. notilla 
were made at the river-bottom; here, studies were confined to a strip 
of very dense forest understory bordering the bank of the Rio Sarapiqui. 
The same area of river-bottom forest has been the study site for similar 
studies on the biology of Itaballia caesia (Pieridae) (Young, 1972c), 
the ithomiine Hymenitis nero (Young, 1972a), and the nymphalid 
Victorina epaphus (Young, 1972e). This area was visited a total of 10 

VoLUME 28, NUMBER 3 259 

days during July and August of 1971 for the sole purpose of studying 
P. notilla. Usually no more than 3 to 4 hours during the morning were 
spent here each day. 

During July 1972, we found a thin strip of clearing that ran up the 
slope of the ravine at a point further west of this river-bottom site, and 
very close to a second wooden bridge (the one not having a waterfall 
near it) at a hairpin turn in the road to Puerto Viejo. The clearing 
was made by the I. C. E. (Instituto Costarricense de Electricidad) during 
the installation of a telegraph line across the Rio Sarapiqui; the vegetation 
under the line is cut down at least twice annually. Here we searched a 
total of five days for eggs, larvae, and foodplants of P. notilla, within the 
dense understory immediately to either side of this strip, and within the 
thinned-out vegetation of the strip itself. We worked a distance of about 
100 m, from river-bottom to the road on top, spending about three hours 
each day doing only this work. Although the butterfly was seen fre- 
quently in the second-growth along the roadcut near the ridge-top, we 
did not make any attempts to study it there. 

Field studies of P. notilla included observations of habitat selection by 
flying adults, observations on oviposition behavior, determinations of 
larval foodplants, and note-taking on larval behavior. All of these studies 
were conducted each day we visited the river-bottom at Cuesta Angel. 

Laboratory studies consisted of describing life stages and estimating 
mean developmental time from egg to adult. The “laboratory” was a 
converted tool shed on the premises of the Costa Rican program of the 
Associated Colleges of the Midwest in San Jose, Costa Rica. Eggs were 
collected in the field at Cuesta Angel and transported by jeep to San 
Jose within one or two days. The eggs were confined to clear plastic 
bags (each one 8 X 20 cm) containing fresh cuttings of the foodplant. 
We inspected immatures every one to three days, measuring body length 
of larvae, collecting head capsules, and examining color patterns. A 
total of 25 eggs were collected for these studies, all within a three day 
period, and divided into five laboratory cultures each containing five 
eggs. The 25 eggs represent a total of seven oviposition sequences in 
the river-bottom study area. Probably several different females were in- 
volved in the egg-laying, so that genetic differences may be a source of 
variability in estimating developmental time. 

Laboratory conditions were 21-23° C and 40-60% humidity for the 
30-day rearing period in San Jose. The cultures were kept on a table 
in a shaded part of the shed. Foodplant was replenished every 3-4 days 
and bags were wiped clean of excess moisture and feces. The same 
techniques have proven successful for rearing immature stages of several 


different groups of tropical butterflies with minimal mortality (Young, 
1972a, b, c, d, e, f; Young & Muyshondt, 1972, 1973). 


Habitat and larval foodplant. The butterfly (Fig. 2,A) is found 
throughout the slopes of the ravine, but adults are more abundant in 
shaded forest understory, especially where it borders thinned-out areas 
of second-growth. The uniform abundance of the butterfly at various 
points on the side of the ravine indicates that the species is not responding 
to any gradients in micro-environmental factors. But a difference in 
larval foodplants exists between the river-bottom and higher places within 
forest understory on the side of the ravine: the single larval foodplant 
found at the river-bottom is Cestrum megalophyllum Dum. in the 
Solanaceae. Here, the plant occurs as a small woody understory tree 
that grows to about 3 m in height. The tree can often be found growing 
in small groups of 2-5 individuals, although these groups are patchily- 
distributed in the understory. The uppersides of the very large con- 
spicuous leaves of this species are often covered with moss and other 
forms of epiphytic growth. 

Further up the side of the ravine, another larval foodplant is an 
unidentified species of Capsicum, also a member of the Solanaceae. 
This species represents another small woody member of the understory. 
But along the cleared strip of vegetation made by the I.C.E., there are 
extensive growths of suckers from the cut-down trunks of the original 
trees. The leaves of these suckers are generally larger than those of the 
original trees and often much lighter green in color. Only these two 
solanaceous species at Cuesta Angel provide oviposition sites and larval 
food for P. notilla. 

Life cycle and developmental time. The oblong-shaped egg is 1.2 
mm high by 1.0 mm wide at the middle. It is marked by several vertical 
grooves, and the top is rounded (Fig. 2,B); the egg is uniformly white 
until the hatching of the first instar larva. 

The first instar larva is generally dark green in color, once it begins 
to feed on plant tissue. By the time of the first molt, it is about 3.5 
mm long. The first, second, and third instars are virtually identical in 
appearance (Fig. 2C,D,E). Each larva is dark green dorsally. On 
each side dorsolaterally, a thick light green line runs from the first 
thoracic segment to the anal plate (Fig. 2,E). Beneath this pair of lines 
the body continues to be dark green for an additional fraction of a 
mm; then this color gives away to light translucent green. The head in 


Fig. 2. Life stages of Pteronymia notilla: (A) adult (dorsal and ventral aspects); 
(B) egg; (C) second instar, dorsal aspect; (D) second instar in curled-up position 
(presumably defensive); (E) third instar, lateral aspect; (F) fourth instar, lateral 
aspect; (G) fifth instar in curled-up position (presumably defensive); (H) fifth 
instar, dorsal aspect; and (I) pupa, lateral aspect. Dimensions of life stages are 

given in the text. 


all three of these instars is shiny black, but has a mask-like appearance, 
resulting from a three-pronged, forked light green line, that superficially 
divides the head into three regions (Fig. 2,E). The anterior edge of the 
first thoracic segment behind the head is enlarged and orange-yellow. 
The anal plate is dark green and bordered with thick patches of yellow. 
The true legs are dark green and the false feet are light green. The 
second instar attains a length of 6.5 mm by the second molt, and the 
third instar is about 12.0 mm long by the third molt. 

The fourth and fifth instars are identical to one another in coloration, 
but very different from the previous three instars (Fig. 2F,G,H). The 
dorsal color pattern consists of a thin medial light blue line running from 
the first thoracic segment to the anal plate, and bordered to either side 
by an alternating series of short light blue and dark green bands, 
running perpendicular to the central blue line (Fig. 2,H). These series 
of bands do not extend to the head and anal plate: anteriorly, there is 
a swollen region just behind the head, and posteriorly, there is another 
one just before the anal plate. The anterior swollen region forms a light 
green collar ringed with orange; the posterior swollen region is uniformly 
bright orange, but does not cover the entire dorsal region (Fig. 2,H). 
The light green thick dorso-lateral line of the previous instars is now 
yellow, and the body beneath it is light green. The thin, central blue 
line continues through both swollen areas. Just behind the swollen anal 
region, there is one segment bearing the typical body color pattern. 

The head of the fourth and fifth instars now appears to be much 
smaller due to the swollen aspect of the anterior trunk segments. It is 
shiny black with the inverted “Y” portion of the light green line pattern 
being thicker than in the previous instars (Fig. 2,G). Finally, there is 
a thin yellow lateral line running the length of the body, located just 
where the ventrum joins the lateral aspects of the body. The fourth 
and fifth instars are much more brightly colored than the previous in- 
stars. The larva is about 16 mm long by the end of the fourth instar, and 
about 22 mm long by the end of the fifth instar. 

Immediately prior to pupation, the fifth instar larva contracts in length 
and becomes a uniform green color. This prepupa produces a pupa ( Fig. 
2,1) which is uniformly light green and slightly reflective. The pupa is 
remarkably translucent with only abdominal regions being clouded over 
with a yellowish coloration just beneath the cuticle. The cremaster is 
light red. The pupa is about 17 mm long by 7 mm wide (dorsoventrally ) 
through the thoracic region. The coloration of the pupa does not change 
appreciably prior to the eclosion of the adult. 

There is very little sexual dimorphism in coloration of the wings in 

VoLUME 28, NUMBER 3 263 

TaBLE 1. The developmental time (days) of Pteronymia notilla on Cestrum 
megalophyllum (Solanaceae) in the laboratory.* 

ae ee ee ee ore ee ee ee EGG- 
EGG 1 ® 3 4 5 PUPA ADULT 
MEAN 5 2 D) 3 5 6 7 30 
a2 Side SE (U3 “se (Q.Y ae Mil sel se Quo sey Se abil 
No. Individuals 
Measured (N) 94 D4 it malt all 21 20 

* All measurements were made in one laboratory in San Jose, Costa Rica. During this time, 
laboratory conditions were 21—23° C and 40-60% relative humidity. See text for further details. 

the adult (Fig. 2,A); good descriptions are given by R. Haensch in Seitz 
(1924) and by Fox (1968). For a total of 20 individuals reared in the 
laboratory, the mean length of the forewing is 25 + 0.7 mm, which is 
very similar to forewing length of wild caught individuals. 

The egg through adult developmental time in the laboratory required 
30 days (Table 1). Developmental time is undoubtedly quite variable 
in the field. 

Larval behavior. The larvae of P. notilla generally occur singly on 
leaves of the foodplant; there is no evidence of gregarious behavior when 
more than one larva is present on an individual plant. Both resting and 
feeding are confined to the undersurfaces of leaves, and the larvae of 
all instars are most frequently found in the field on older leaves. Pupa- 
tion often occurs on the foodplant and both living pupae and hatched 
pupal cases have been found on the undersurfaces of leaves attached 
along a major rib. The earlier instars (1-3) are very cryptic in ap- 
pearance, and are very difficult to find on foodplants in the wild. 
Despite the increased conspicuousness of the later instars (4-5), there 
are no noticeable changes in larval habits and behavior. Individuals of 
all instars exhibit a pronounced curling up behavior upon tactile contact 
with forceps (Fig. 2D,H); this behavior may be defensive. 

Individual larvae build silken trails over leaves and stems, but there 
is no nest construction as seen in the solitary larvae of Hymenitis nero 
(Young, 1972a). Furthermore, there is no “dropping off” behavior, where 
individual larvae suspend themselves from long silken threads as a 
means of escaping predatory attack. Such behavior has been noted for 
various ithomiine larvae, and it has recently been seen in Dircenna relata 
where the larvae are semi-gregarious (Young, 1972b). 

Adult behavior. Adults are often seen flying about 1-2 m from the 
ground in forest understory. Presumably adults spend a substantial 


amount of time cruising for courtship encounters and searching for 
Oviposition sites. Courtship has not been observed in P. notilla. The 
reproductive strategy of P. notilla involves carefully laying each egg 
singly on the ventral surface of older leaves of Cestrum megalophyllum 
and Capsicum sp. Eggs are also laid on the large leaves coming from 
suckers of cut-down trees. 

In a total of 13 oviposition sequences observed on five different dates, 
there were six in which the female laid more than one egg on a single 
leaf. In these instances, there were no more than three eggs laid on 
the leaf. Furthermore, the eggs were never close to one another, but were 
widely scattered on the under surface of the leaf. On a visit to a single 
foodplant tree, an ovipositing female would lay anywhere from one to 
seven eggs in the tree; there were never more than three eggs on a leaf 
when multiple ovipositions were seen. 

Egg laying involved the female landing on the ventral surface. The 
female walked toward the interior of the leaf and laid the egg. If 
more than one egg was to be laid, there were brief periods of walking 
before laying the next egg. Eggs were never laid near the edges of the 
leaves. Oviposition has been observed at various times throughout the 
morning, but seldom during the afternoon hours. There appears to 
be no correlation between time of oviposition and the amount of sun- 
shine filtering down through the forest canopy. Females often rest 
for several minutes between oviposition sequences. 


P. notilla differs from other ithomiids in a number of ways: first, 
egg color and external morphology when compared with that of two 
other recently studied species in Costa Rica, namely, Hymenitis nero 
and Dircenna relata (Young, 1972a,b). The egg of H. nero is white 
and less oblong than the egg of P. notilla, but it has the same distribution 
of vertical grooves as in the latter. The egg of D. relata is deep yellow 
and has the general shape of the egg of H. nero; but unlike both this 
species and P. notilla, there is a complex series of short horizontal grooves 
evenly-spaced between adjacent vertical grooves on the external surface. 

The larval stages of these three species are very different in appearance 
and of these three, only the fourth and fifth instars of P. notilla show a 
dramatic change in coloration from the previous instars (Fig. 2); similar 
changes in color are not seen in the other two species. The larvae of 
both H. nero and D. relata retain a generally mottled green cryptic ap- 
pearance throughout all instars (Young, 1972a,b). Furthermore, the 

VoLUME 28, NUMBER 3 265 

pupa of D. relata is extensively covered with gold coloration, especially 
on wing pads and dorsal aspects of the thoracic and abdominal regions. 

The pupa of H. nero is heavily adorned with a bright silver coloration 
on the wing pads and thorax. Such highly reflective silver or gold 
pigmentation is entirely absent from the pupa of P. notilla, in which 
protective coloration is limited to light green translucence with rela- 
tively minor reflectance properties. Finally, the developmental time for 
these three ithomiid butterflies is between 25 and 30 days in the 

These three species also illustrate ecological divergence in the 
Ithomiidae with respect to larval foodplants. While it is known that 
most Ithomiidae feed on various Solanaceae (e.g. Brower & Brower, 1964; 
Ehrlich & Raven, 1965), little is known about patterns of divergence in 
foodplant exploitation at the generic and species levels among these but- 
terflies. Such information is clearly of great importance in studying the 
community structure of the butterflies. 

In the present situation, at least two sympatric ithomiids, P. notilla 
and H. nero, exploit different species of Cestrum in the forest understory 
of Cuesta Angel. D. relata is found in S. hispidum at one locality 
(Bajo la Hondura) on the Pacific slopes of the Central Cordillera (Young, 
1972b); the butterfly also occurs at Cuesta Angel, but the larval food- 
plant has not yet been determined. However it is likely that this species 
feeds on different foodplants than both P. notilla and H. nero at Cuesta 
Angel. While H. nero is a blue clear-winged species of ithomiid, P. 
notilla is one of the more conspicuous orange-winged species. The showy 
coloration of the late instars of the latter species may be indicative of 
noxious or unpalatable properties of older larvae, pupae, and adults. 
Clear-winged species such as H. nero are presumably more palatable, 
since their immature stages employ a more pronounced strategy of crypsis 
than is seen in P. notilla: the larvae are cryptically-colored throughout 
all instars; the pupae are more effective in resembling large drops of 
rain water hanging from leaves (e.g., Brower, 1971); and the larvae 
construct nests of partially closed leaves where they rest when not 
feeding (Young, 1972a). 

Furthermore, the transparent qualities of the wings make the adults 
rather inconspicuous in shaded forest understory. This apparent di- 
vergence in adaptive strategy is interesting since both species feed on 
related species of Cestrum. Such a divergence in larval feeding habits 
is illustrative of very subtle environmental factors (i.e., species differences 
in secondary compounds among congeneric sympatric plants), which 
influence the evolution of morphological and behavioral traits among 


herbivorous larvae toward either crypsis or warning coloration, two very 
different adaptive strategies. 

But orange-winged species of ithomiids, like P. notilla, may not be 
as unpalatable as other orange-winged genera such as Dircenna and 
Mechanitis. This is suggested by several factors: (1) oviposition in 
D. relata is semi-clustered, while it is single in P. notilla (Young, 1972b); 
(2) oviposition is clustered in M. isthmia; (3) larvae are semi-gregarious 
in D. relata (Young, 1972b) and gregarious in M. isthmia, but solitary 
in P. notilla; (4) the dorsal wing surfaces of D. relata are brighter than 
those of P. notilla, whereas those of M. isthmia are strongly mimetic, 
since they have familiar tiger-striped pattern of various heliconiids and 
the danaid Lycorea. 

The observed differences in life cycles, larval foodplant utilization, 
and dorsal wing surface coloration among different genera of the 
Ithomiidae suggest that differences in adaptive strategy with respect to 
escape from predators have evolved. One lesson to be learned from such 
preliminary assays of ithomiine natural history is that experimental 
feeding studies utilizing a wide range of vertebrate and invertebrate 
predators must be performed to demonstrate differences in the relative 
palability of adults and larvae among different genera. Such studies 
must be accompanied by field studies elucidating various behavioral 
patterns (e.g., communal roosting, alarm positions, etc.) which may be 
correlated with increasing unpalatability in heliconiid butterflies ( Benson, 


(1) The life cycle and developmental time of the ithomiid Pteronymia 
notilla Butler & Druce are given for individuals reared from eggs col- 
lected at one montane tropical forest locality in central Costa Rica. The 
developmental time in the laboratory is about 30 days and fourth and 
fifth instar are brightly colored relative to earlier instars. 

(2) The major larval foodplant at the bottom of the ravine where 
the species was studied is Cestrum megalophyllum (Solanaceae). Further 
up the side of the ravine, another foodplant is Capsicum sp. (Solanaceae). 
Both species occur as small woody understory trees. 

(3) Both eggs and larvae generally occur singly on the foodplants, 
and there is no evidence of cluster oviposition and larval gregariousness, 
as noted in other ithomiids. | 

(4) Oviposition is precise in this species and involves the female 
walking to a suitable spot on the ventral leaf surface before an egg is 

VoLUME 28, NUMBER 3 267 

laid. Females seem to show some selectivity, preferring to oviposit on 
older leaves. 

(5) The noticeable change in larval appearance at the third molt 
is suggestive of increased unpalatability, which may be carried over to 
the adult stage. Orange-winged ithomiids such as P. notilla appear to 
the human observer more conspicuous than clear-winged species of com- 
parable wingspan. The unpalatability of dull orange species like P. 
notilla, however, may be weak, since some of the more bright-orange 
genera (Dircenna and Mechanitis) have life cycles in which oviposition 
is clustered and larvae are gregarious. These forms are presumably more 
unpalatable than similar appearing ithomiids with solitary oviposition 
habits and non-gregarious larvae. Such correlations, however, are very 
tentative, in the absence of experimental data on the relative palatability 
of adults and immatures for representatives of different genera. 


I am very grateful to Lawrence University for supporting this research 
through a College Science Improvement Grant (COSIP-GY-4711) during 
the summer of 1971. Laboratory facilities and logistic support was pro- 
vided by the Costa Rican Field Studies Program of the Associated 
Colleges of the Midwest. Patrick Eagan assisted in all aspects of the 
field and laboratory work. Drs. Lee D. Miller and Keith S. Brown, Jr. 
identified the species studied. Dr. Dieter C. Wasshausen of the Smith- 
sonian Institution identified the larval foodplants. 


Benson, W. W. 1971. Evidence for the evolution of unpalatability through kin 
selection in the Heliconiinae (Lepidoptera). Amer. Natur. 105: 213-226. 
Brower, L. P. 1971. Prey coloration and predator behavior. In Topics in the 
Study of Life, Biol. Source Book, Sect. 6, Anim. Behav. Harper & Row, N.Y. 
p. 300-370. 

& J. V. Z. Brower. 1964. Birds, butterflies, and plant poisons: a study in 
ecological chemistry. Zoologica 49: 137-159. 

EuecuicuH, P. R. & P. H. Raven. 1965. Butterflies and plants: a study in coevolu- 
tion. Evolution 18: 586-608. 

Fox, R. M. 1968. Ithomiidae of Central America (Lepidoptera: Nymphalidae). 
Trans. Amer. Entomol. Soc. 94: 155-208. 

Serrz, A. (ed.) 1924. Macrolepidoptera of the World. Vol. 5. The American 
Rhopalocera. Verlag, Stuttgart. 

Younc, A. M. 1972. On the life cycle and natural history of Hymenitis nero 
(Lepidoptera: Ithomiinae) in Costa Rica. Psyche 79: 284-294. 

1973. The life cycle of Dircenna relata (Ithomiidae) in Costa Rica. J. 

Lepid. Soc. 27: 258-267. 

1972b. A contribution to the biology of Itaballia caesia (Pierinae) in a 

Costa Rican mountain ravine. Wasmann J. Biol. 30: 43-70. 


. 1972c. Breeding success and survivorship in some tropical butterflies. 
Oikos 23: 318-326. 

1972d. The ecology and ethology of the tropical nymphaline butterfly, 
Victorina epaphus. I. Life cycle and natural history. J. Lepid. Soc. 26: 155-— 

. 1973b. Notes on the biology of the butterfly, Heliconius cydno ( Lepidoptera: 
Heliconiinae ) in Costa Rica. Wasmann J. Biol. 31: 337-350. 

& A. MuysHuonpr. 1972. Biology of Morpho polyphemus (Lepidoptera: 
Morphidae) in El Salvador. J. N. Y. Entomol. Soc. 80: 18-42. 

& A. Muysnonpr. 1973. Notes on the biology of Morpho peleides in Central 
America. Carib. J. Sci. 13: 1-49. 


Recently Clench (1972, Ann. Carnegie Mus. 44: 33-44) described a new species 
of Lycaenidae, Celastrina ebenina. This butterfly was formerly known as a “black 
form” of the common C. argiolus pseudargiolus: form ¢ nig and form @ intermedia 
as listed by dos Passos (1964, Lepid. Soc., Mem. 1: 69, 481). Clench asked me 
to be on the lookout for this species and on 29 April 1972, I took two males in 
Buncombe County, North Carolina, and sent them to him. My find extended 
the confirmed range into North Carolina. 

On 21 April 1973 I took another male and on 4 May 1973 I found a single 
female. All of the ebenina I have taken were found in Buncombe County, North 
Carolina, along the dirt road which is an extention of Buncombe County road number 
2178 south of its junction with county road 2173 at Dillingham, a small community 
near Barnardsville. The two taken in 1973 were found about 2.1 miles south of the 
junction near the parking place on the left side of the road (elevation about 2880’). 
A mountain stream parallels the road on the right at this point. The two taken in 
1972 were found about 3 miles south of the junction (elevation about 3260’). 
This road runs from Dillingham to the Blue Ridge Parkway, and the locations 
described can therefore be reached by driving north (down the mountain) from 
the Parkway. 

I am publishing this note to encourage other collectors to look for this butterfly 
in the southeastern mountains of the United States. According to Clench it should 
be sought in cool, moist, forested ravines and is almost always found near areas 
where Trillium grandiflorum is in bloom. The habitat in which I took ebenina matches 
perfectly with this description which Clench gave of the other areas in which it 
has been taken. I would be pleased to hear from others who find it. 

RicHARD E. Price, Jr., P.O. Box 146, Mars Hill, North Carolina 28754. 

VOLUME 28, NUMBER 3 269 



A.R.C. Unit of Invertebrate Chemistry & Physiology, 
Department of Zoology, Downing Street, Cambridge, England 

In about 1860 Pieris rapae L. (the imported cabbage worm) was 
recorded from Canada (Seitz, 1924). It spread rapidly and already 
by 1870 was causing great damage to cruciferous truck crops from 
Montreal to New York and within a surprisingly short time had spread 
throughout the Union (Chittenden, 1905). 

In 1972 I received for identification some white butterflies from the 
region of Santiago, Chile. They are undoubtedly Pieris brassicae L., 
the large cabbage white butterfly, to give it its English vernacular name. 
The specimens were forwarded to me by fellow member J. H. Robert 
who had received them from Sr. Luis E. Pefia who reports that they 
are now (1972) “flying around gardens in the vicinity of Santiago” 
which means the species is clearly established and was doubtless 
introduced some years ago. In view of the enormously rapid rate 
of spread of which Pieris species are so clearly capable it would seem 
desirable to give some details of it so that it can be immediately 
recognised and dealt with, if that be possible. Already grave concern 
is being expressed about an African honeybee, Apis mellifera adansonii, 
which is sixteen years has spread virtually throughout the whole of 
South America and is heading fast toward the U.S. (Orsak, 1973). It 
would seem quite possible for P. brassicae to follow the same course, 
it is a noted migrant and just as fond of Cruciferae as is P. rapae; indeed 
its larvae will feed on plants of any family containing mustard oil 
glucosides. However, in the Canary Islands it is not a pest, the larvae 
feeding only on Tropaeoleum (Fernandez, 1955). 

The probability that it will now spread through South America 
appears to be a very real one, as Sr. Pefa informed me in June 1973 
that it is already widespread in all the Province of Valparaiso and adults 
are already flying in other provinces and the Cruciferae are being 
destroyed. It would seem desirable therefore to take the opportunity 
to give some account of the species, so that it can be looked out for; 
and at the same time to correct certain errors concerning it in the 
literature and put on record some new observations. P. brassicae differs 


Fig. 1. Typical eggbatch of P. brassicae. 

quite markedly from other European members of the genus. There 
are some quite good reasons for considering that it should be separated 
off into another genus, and it is only the great confusion that this would 
cause that seems to have prevented this step from being irrevocably 

Ege. Fig. 1 shows a batch of eggs and Fig. 2 eggs in situ on cabbage. 
They are laid in more or less regularly arranged batches which vary 
in size from a few eggs to a hundred or more, the number varying 
according to the age of the butterfly, with an average around 40-50. 
When first laid the eggs are a very pale straw color; within twenty four 
hours this has darkened to yellow and in at least one subspecies (P. b. 
cheiranthi Hueb) they are bright orange. Eggs from butterflies whose 
larvae have been reared on semi-synthetic diets not containing cabbage 
leaf powder, remain a very pale straw color, indeed may be almost 
white. A female is capable of producing 750 eggs during a full lifespan 
(David & Gardiner, 1962) but it is doubtful if the full number is ever 
produced under feral conditions. In very warm weather the eggs will 
hatch in 4-6 days but may well take 2-3 weeks in cold weather. A 
few hours before hatching the eggs turn black and the form of the 
larva can be seen through the shell. The first larvae to hatch turn 
round and often commence to eat the tops of the shells of the other 


larvae. In this fashion the hatch of a batch of eggs will take place 
over about 30 minutes. The young larvae consume the eggshells and 
then, en mass, spin a silken pad on which they rest when not feeding. 

Larva. Fullgrown larvae are shown in Fig. 3A. The larvae are 
gregarious throughout their life, unlike P. rapae and other “small white 
species’ which are not only solitary, but cannabalistic, both to smaller 
brethren and, in particular, to their own eggs, which they eat and 
kill (unlike brassicae which merely eat the top of the shell and release 
the contained larva). They invariably have five instars and, depending 
on the temperature, the larval stage lasts from two to eight weeks. 
It has been erroneously stated by both Klots (1958) and more recently 
again by Wigglesworth (1972) that five instars only occur under cold 
conditions and that the number of instars falls to four and finally 
only three as the temperature of rearing increases. These statements 
are based on an observation of Klein (1932). Frohawk (1934), a 
careful recorder, who had the experience of rearing every species of 
British butterfly, considered only five instars, and David & Gardiner 
(1962a) proved conclusively that the number of instars is constant at 
five over the very wide range of environmental conditions at which 
rearing is possible, and further extensive rearing by the present author 
with various stocks and races of brassicae has subsequently confirmed 
this. The color of the larvae is virtually the same in all instars; blue- 
grey or yellowish ground color, a yellow dorsal stripe and irregular 
and intricate black markings which are more intense the lower the 
temperature of development. The yellowish ground color is recessive to 
the blue-grey (David & Gardiner, 1962a) but appears to be so common 
in the wild that there must be some advantage in it. The larvae prefer 
to feed openly on the outside of the leaves. Fig. 3B shows an aggregate 
of mainly fourth instar larvae and Fig. 4 the remains of a garden cabbage 

Chrysalis. These are formed in a similar fashion to those of P. 
rapae, that is to say suspended by a cremaster and a silken girdle. Also 
as in P. rapae similar situations are sought by the larvae in which to 
pupate. The color of the chrysalis is either a pale straw or a shade of 
green, with variable black markings, and in general the color is lighter 
or darker according to the background. It has been stated (Babers & 
Pratt, 1952) that the color is influenced by the illumination of the larva 
before pupation. As a result of numerous experiments and the rearing 
of more than one million larvae I have never found any evidence of 
this. I have, however, found conclusive evidence that diapausing 
chrysalids are much more inclined to be green in color than summer 


Figs. 2-4. P. brassicae: 2, eggs in situ on cabbage leaf; 3A, fullgrown larvae 
on same plant; 3B, gregarious cluster of mainly fourth instar larvae; 4, devastated 
crop of cabbages in a garden plot. 

brood ones (Gardiner, in prep.). This was strikingly born out by the 
chrysalids I received from Chile, the straw-colored ones eclosed a few 
days after receipt; the green-colored ones are still unchanged after 
several weeks and therefore clearly in diapause. 

The pupal stage of summer brood specimens lasts 10 days in warm 
weather, but may be as long as 60 days if the weather is cold. If the 
pupa has entered diapause then this stage will last for 6-8 months. 

Adult. In general appearance the adults of P. brassicae are similar 
to those of the imported cabbage worm P. rapae, but are quite distinctive 
and sexually dimorphic. In particular the black markings have a sharp 
cut-off from the white instead of the gradual fade-out from one to the 
other as in P. rapae and P. napi. Both sexes are white with a black apical 
spot. The female only, has two black distal spots and a black discal 
streak along the inner margin. Both sexes have two black discal spots 
on the underside. 

The underside of the hindwing tends to be very variable and may 
be yellow to orange (race cheiranthi); or pale straw, greenish, and at 
times almost black. Greenish and blackish forms are an over-all effect 

VoLUME 28, NUMBER 3 Dits 

produced by a light to heavy sprinkling of black scales. Unlike other 
Pieris species the veins of the wings in brassicae are never heavily marked 
to give a rayed or chequered effect. In size brassicae is larger than all 
other United States Pieris with a wingspan of from 55-65 mm in certain 
bred examples (David & Gardiner, 1961), up to 63-76 mm in wild 
caught specimens (Frohawk, 1934). All other United States Pieris have 
a wingspan of under 50 mm (Chang, 1963). 

P. rapae crucivora from Japan are exceptionally large. Esaki & 
Yokoyama (1955) give the wingspan as 55 mm and I have bred specimens 
up to 60 mm. Fig. 5 shows one such bred specimen for comparison 
with brassicae. Since Esaki & Yokoyama use a different basis for their 
wingspan measurement than Chang (which gives a lesser figure), the 
actual size of P. r. crucivora comes out as the mean of the P. brassicae 
bred by David & Gardiner (1961). As in other Pieris the black coloring 
of the spring brood is much paler than in the summer broods. 

The Chilean examples have the typical upperside facies, but the 
hindwing underside is of the dark green form. This form certainly 
occurs in British, Spanish, German, and Maltese race wollastoni, and 
in East European brassicae, but an examination of my collection and 
of the extensive series in the insect room of the Cambridge University 
Museum of Zoology reveal that it is uncommon, the lighter forms being 
by far the more numerous. It does not therefore seem possible to pin 
down the exact origin of the Chilean P. brassicae. It has been suggested 
by Sr. Pena that they may have come from Eastern Europe, there now 
being considerable trade between there and Chile. What is more 
certain is that the specimens are not of one of the numerous races of 
P. brassicae, which has distinct forms in certain parts of its range where 
it also appears to be non-migratory. Details of the distribution are 
given in Fig. 13. 

Three of the Chilean specimens are shown in Figs. 6,7,8. For com- 
parison, a typical English pair (Figs. 9,10), the male English underside 
(Fig. 11), and the dark green form of a Spanish example (Fig. 12) are 

The dark green color on the hindwing underside of the Chilean 
examples is interesting. This may well be due to adaptation to certain 
environmental factors which confers some advantage in a particular 
area. It has already been shown by Gardiner (1973) that the facies of 
brassicae can be changed by careful selective breeding, and the dark 
green and the yellow form of the hindwing underside are amongst 
the characters which can be so selected. The Cambridge stock of 
brassicae, as was maintained for so many years by David & Gardiner, 


Figs. 5-12. Various imagoes for comparison: 5, P. rapae crucivora Q ex Japan; 
6-12 P. brassicae: 6, 6 underside ex Chili; 7, 2 upperside ex Chile; 8, ¢ under- 
side ex Chili; 9, 2 upperside ex David & Gardiner’s “Cambridge” stock; 10, ¢ 
upperside ex “Cambridge” stock; 11, ¢ wumderside ex “Cambridge” stock; 12, ¢ 
underside ex Spain. 

VoLUME 28, NUMBER 3 Pals) 


Fig. 13. Palearctic distribution of P. brassicae (within heavy line), and area 
in South America from which now recorded. 

has rather a light straw-colored underside and I have similar specimens 
in my collection from most areas of Europe and also from the North 
African litoral and near East. It will be interesting to hear in due course 
if all Chilean examples are of this dark green form or if the lighter colored 
ones are also to be found. The question of this underside coloration 
presents a good opportunity for some field research. Eastern European 
and Asiatic material is not so readily available but I have seen all types 
of underside from those areas. 

P. brassicae is a well known migrant. Although, due to destruction 
of former breeding areas, very vast swarms no longer occur, regular 
migration usually in a southerly and westerly direction still takes place. 
Return flight does not occur. The insects migrate within a day or two 
of emergence, the females often mated, but not yet with mature eggs, 
and are capable of traversing up to 250 miles, without food, in a few 
days. (For further details, see Johnson, 1969.) It can therefore readily 
be appreciated that, once a nucleus colony is established, a very rapid 
spread of the butterfly can take place. 

Diapause. P. brassicae has a facultative diapause controlled by the 
daylength on the larva (Way, Smith & Hopkins, 1949; David & Gardiner, 
1962a). Consequently as long as the daylength is sixteen hours or longer, 
dawn to dusk, and the temperature averages above 10° C, there will 
be a continuous succession of broods, at least one every six weeks in 
very warm tropical weather. As soon as the daylength falls the pupae 
will enter diapause. All summer brood stages can withstand frost for 


short periods and diapausing pupae can withstand severe and prolonged 
winter conditions. There is no doubt that the species must be considered 
very hardy. 

Parasites, predators and diseases. P. brassicae larvae are parasitized 
by a number of Apanteles species, in particular A. glomeratus L., which 
is known to attack P. rapae in the United States (Blunck, 1957). The 
pupa is also parasitised by Pteromalus puparum L., which is believed 
to have been imported into the States at the same time as P. rapae which 
it also attacks (Chittenden, 1905). Both larvae and adults are also 
predated by social Hymenoptera. The only avian predator which has 
been observed eating the very distasteful larvae is the European thrush, 
Turdus musicus L. Flying adults are sometimes attacked but appear 
to be rarely eaten by birds, although mice (Mus musculus L.) will eat 
the bodies. The pupae, however, are eaten in considerable numbers 
(Moss, 1933). Eggs do not seem to be attacked by any parasite or 
predator and even larvae of its own kind have been observed by me 
to eat carefully round eggbatches without doing them any damage. 
Various potential parasites already present in the Nearctic region, notably 
A. rubecula, which helps to control P. rapae and could also attack P. 
brassicae, have recently been surveyed by Blunck (1957) and Wilkinson 
(1966). Various species of ant have been observed in England to carry 
off and consume the young larvae. Microsporidian parasites are recorded 
from Europe (but not England), but Blunck (1957) could not find 
these in the United States. 

P. brassicae is certainly susceptible to many of the usual commercial 
insecticides and also to Bacillus thuringiensis. From time to time the 
larvae and pupae succumb to an undescribed bacteria, but it is my 
experience that they are far less susceptible in this respect than many 
other species of Lepidoptera. However, they are very susceptible indeed 
to a granulosis virus disease. Although the virus might be the better 
method of control, it is not yet commercially available, although B. 
thuringiensis is. (For data on these two possible control agents, see 
Burges & Hussey, 1971.) Although certainly susceptible to many in- 
secticides, brassicae is difficult to eradicate and its present-day cessation 
as a major pest in large parts of the palearctic region is due in my view 
not so much to control measures as such, but to changed agricultural 
practices and, above all, to the bringing into other uses of enormous 
areas of its former wild breeding areas with consequent wholesale 
destruction of its foodplants in these areas. The enormous migrations 
recorded fifty and more years ago no longer occur. 



I am indebted to Sr. J. H. Robert of Alicante, Spain for forwarding 
to me the original Chilean specimens and for Spanish examples of P. 
brassicae; to Sr. Luis E. Pefia of Santiago, Chili for subsequent live 
material and information from Chili; to Hr. Hermann Wilde of 
Darmstadt, Germany for Fig. 4 and to Mr. G. H. Runnalls and Miss 
Yvonne R. Carter of this Department for photographic help. 


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11: 129-140. 

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WiIcGLESworTH, V. B. 1972. The Principles of Insect Physiology. Chapman & Hall, 

Wixkinson, A. T. S. 1966. Apanteles rubecula, Marsh, and other parasites of Pieris 
rapae in British Columbia. J. Econ. Entomol. 59: 1012-1013. 




Ukrainian Research Institute for Plant Protection, 
33 Vasilkovskaya Street, Kiev 127, Ukraine 252627, U.S.S.R. 

A study of the composition, spatial differentiation and zoogeographic 
connections of the Caucasus butterfly fauna requires verification of the 
species determinations given by former authors. Direct comparison of 
material collected in the Caucasus area with published data often shows 
disagreement between the species and the names attributed to them 
by different authors. Such a case has been exemplified recently with 
Eumedonia eumedon Esper (Nekrutenko, 1972). One of the most 
important points in taxonomically-based faunistic speculations is good 
knowledge of the fauna in adjoining areas. It would be no exaggeration 
to say that a reliable, ‘three dimensional’ picture of the Caucasus but- 
terfly fauna requires two essential conditions: familiarity with the 
European fauna in order to compare the local forms with their 
nomenotypic subspecies and, on the other hand, knowledge of the fauna 
of Turkey and Iran in order to detect clinal intergradations where they 

In this paper I consider another case of taxonomic uncertainty re- 
garding the position of the Caucasian representative of the [sub] genus 
Agriades Hiibner (Polyommatus Latreille, pars), and I describe a new 
subspecies of A. pyrenaicus from Ulu-Dag, Turkey, as a link in the 
intergrading chain of its geographic subspecies. 

As is fairly obvious from synonymic lists, under the description of 
A. pyrenaicus latedisjunctus Alberti, authors almost unanimously have 
attributed Caucasian Agriades to dardanus Freyer, which is considered 
by them to belong, as a subspecies, to glandon Prunner (= orbitulus 
auct., for history see Hemming, 1967), or to pyrenaicus Boisduval. Such 
a situation necessitates answers to two essential questions: (1) to what 
species does dardanus really belong, and (2) do Caucasian Agriades 
belong to dardanus. As part of the alternative (glandon versus 
pyrenaicus), there are two other possible taxonomic interpretations of 
these forms: (3) to synonymize dardanus with pyrenaicus (Forster, 
1938) and/or (4) to consider dardanus as a distinct species, according 
to Freyer’s (1845) original combination (Sauter, 1968). 

VoLUME 28, NUMBER 3 279 

Figs. 1, 2. Two types of juxta structure in Agriades: 1, glandon and aquilo; 2, 
pyrenaicus and its subspecies. 

As has been shown by Chapman (1908) and Bethune-Baker (1913) 
the peculiarities of the “ancillary appendages,” especially differences in 
the structure of the tip of the upper valval lobe, are of high value for 
recognition, so that there are no problems with exact determination of 
glandon and pyrenaicus on the basis of the male genitalia (see also 
Oberthiir, 1910); however, they are practically unrecognizible on female 
genitalic characters. When dissecting a large sample of both glandon 
and pyrenaicus, collected over an extended area, I found an additional, 
highly exact character permitting the determination of these species at 
a glance with 100% confidence. This diagnostic character consists of a 
pronounced structural difference in the juxta between pyrenaicus and 
other Agriades species, as depicted in Figs. 1 & 2. It is curious that this 
character, so clearly visible on the excellent microphotographs of Chap- 
man (1908), and in illustrations in the recent paper of Fernandez-Rubio 
(1970), was not pointed out in the text of either author and thus seems 
to have been overlooked. 

The type locality of “Lycaena dardanus” was designated by Freyer 
(1845) as “europadische Tirkei.”! The illustrated text of its original 

1 Not “‘Freyer 1844 (Typenfundgebiet ‘“‘Tiirkei”’)” as stated by Alberti (1973). 


Figs. 3-7. The tip of the upper valval lobe (right): 3, glandon, Col d’Allos, 
Basses Alpes, 2500 m, Gallia mer., 11 August 1968, G. Hesselbarth leg.; 4, pyrenaicus, 
Cédre, Htes Pyrénées, Rondou (Zool. Mus. Kiev Univ.); 5, dardanus, Cvrstnica 
Planina, Hercegovina, O. Leonhard leg. (Zool. Mus. Kiev Univ.); 6, latedisjunctus, 
Kazbek Mt., C. Caucasus (Y. Nekrutenko); 7, hesselbarthi, Ulu-Dag, Prov. Bursa, 
Anatolia sept. 17 July 1973, G. Hesselbarth leg. 

description agrees fully with characteristics given by Higgins & Riley 
(1970) of specimens from Cyvrstnica Planina in Hercegovina (Yugo- 
slavia), so that specimens from this locality may be considered as “true” 
dardanus. In addition to specimens from Cvrstnica Planina, in the 
collection of the Kiev State University Zoological Museum, there is also 
a short series of similar specimens labelled “Alibotusch Gebirge, 1900 
m, Al.K.Drenowski leg.,” determined by L. Sheljuzhko (in litt., labels) 
and by Buresh & Tuleshkoy (1930) as dardanus. Dissection of the male 
genitalia showed the complete identity in juxta shape in these two 
samples with pyrenaicus from Pyrenees and latedisjunctus from Caucasus, 
respectively. At the same time, the shape of the upper valval lobe tip 
decidedly differs in glandon, pyrenaicus, “true” dardanus, latedisjunctus 
and hesselbarthi n. subsp. (Figs. 3-7). 

VoLUME 28, NUMBER 3 281 

The aforementioned may lead only to the conclusion that, contrary to 
Higgins & Riley (1970), and in agreement with Bramson (1890), Egorov 
(1903) and Alberti (1970, 1973), dardanus should be considered as 
a subspecies of pyrenaicus, not of glandon. This way, the range of 
pyrenaicus becomes far more extended than is seen from the literature, 
and the occurrence of glandon should be restricted, according to avail- 
able data, to the Alps. However, there are no genitalic characteristics 
to recognize glandon from aquilo Boisduval, a circumpolar holarctic 
species with a significant number of subspecies over its wide range. 
The question of interrelations between these taxa remains open. Also 
remaining open is the question of the possible occurrence of glandon 
(a geographic isolate?) in the Caucasus region. As has been observed 
by Fernandez-Rubio (1970), the spot in the forewing cell (underside ) 
may or may not be present in glandon and its subspecies (e.g. zuellichi 
Hemming). At the same time, this spot is present in all specimens of 
pyrenaicus ever seen in collections or figured in the available literature. 
When counting all names of the specific group involved in Agriades, I 
drew attention to the fact that a specimen of “orbitulus’ araraticus 
Gerhard (Bischoff in litt.) from Turkey, figured and described under 
this (patronymic?) name by Gerhard (1853), showed the lack of this 
spot. This may indicate the conspecificity of araraticus with glandon 
and, thus, the possible occurrence of this species in the Caucasus area. 
However, only a genitalic survey of material available from the eastern- 
most part of Turkey can answer the question of its real taxonomic 
position. Except for araraticus with its uncertain position, all authors 
attribute Agriades of Asiatic Turkey to dardanus (for a review of the 
literature, see Kuznetsov, 1929, p. DLXXII; and De Lattin, 1950). The 
specimens collected in the westernmost part of Asiatic Turkey (Bursa) 
in 1973 by G. Hesselbarth were very different than the ‘true’ dardanus 
and other subspecies of pyrenaicus, and belong to a distinctly marked 
and previously undescribed subspecies. 

Agriades pyrenaicus hesselbarthi Nekrutenko, new subspecies 

(Figs. 8-11) 

General. Smallest butterfly in the group. This subspecies differs from the 
other three hitherto known subspecies of A. pyrenaicus by having no traces of 
the diffused submarginal spots in the hindwing cells Ms-Cu: and M>2-Ms (venation 
and cell terminology after Miller, 1969) in both males and females. Veins do 
not differ by color from the upperside ground color. The underside ground color 
is grey, markings contrasting, almost as in glandon. The female’s upperside is not 
powdered with blue scales. From all other subspecies of pyrenaicus and glandon, 
this one differs clearly by the male genitalia (Fig. 7). 

Male. Length of forewing (base to tip) of holotype 10.7 mm (variation in the 


Sas se ees Be 

_ Figs. 8-11. Agriades pyrenaicus hesselbarthi n. ssp.: 8, 9, ¢ holotype upper and 
undersides, Anatolia sept., prov. Bursa, Ulu-Da& Ms., 2300 m, 17-23 July 1973, 
G. Hesselbarth leg; 10, 11, 2 allotype, upper and undersides, same label data. 

type series 10.4 to 11.0 mm). Upper side of both wings of vivid silvery blue 
color, becoming darker toward the margins. This darker zone begins on the fore- 
wing from discal spot and occupies about 4% of the wing length; between veins 
it does not bear diffused patches of the ground color. At the margins, the dark 
color zone reaches the intensity of the female upperside ground color. Discal spot 
on the upper side of the forewing always contrasting, and because it lies on the 
area shaded with the basal diffused end of the marginal dark zone, it is rounded 
with a bright ring of blue ground color (not white). Fringe white, with black 
strokes at the end of each vein, that do not reach the outer margin. Ground color 
of the forewing underside rather dark, brown. The central cell spot in all specimens 
examined, varied in size, but was always contrasting, rounded with a white ring. 
Postdiscal spots complete, but not as uniform in size and shape as in other sub- 
species, each spot being rounded with a white ring. Submarginal spots complete, 
present in all wing cells. A very narrow, precise dark line goes along the outer 
wing margin. Underside of the hindwing brown, basally powdered with blue 
scales; this bluish zone rather narrow. Black markings rounded with narrow white 
rings. Discal spot with or without black pupil (some dark scales almost always 
present). Yellow submarginal lunule in the cell M;z-Cu: closed with black contrasting 
patches from basal and marginal sides; basally this cell always bears a well developed 
black spot. 

Female. Length of forewing of allotype (base to tip) 10.8 mm (in 3 female 
paratypes ranges from 10.6 to 10.8 mm). Ground color of the upperside of both 
wings dark, brown-black. Black discal spots visible on both wings. Underside color 
and pattern as in male, ground color more vivid, markings developed more strongly. 

VOLUME 28, NUMBER 3 283 

Male genitalia (Fig. 7). General appearance as in all other Agriades. Juxta 
horseshoe-shaped, strongly chitinized. The tip of the upper valval lobe rounded, 
symmetric, head-shaped, bears about 20 teeth. The isthmus between the body 
of valva and the head is well expressed. This character, more than any other, shows 
a similarity to A. pyrenaicus pyrenaicus. Female genitalia. No diagnostic features 
(3 specimens dissected ). 

Material studied. Holotype, male, and allotype, female: Turkey in Asia, Anatolia 
sept., prov. Bursa, Ulu-Dag Ms., 2300 m, 17-23 July 1973, G. Hesselbarth leg. 
Paratypes, 11 ¢ 4, 3 29, same locality, dates and collector. Holotype, allotype 
and 5 646, 2 2@ paratypes and genitalic slides deposited in the collection of 
the Kiev State University Zoological Museum. About 85 paratypes are in the 
collection of G. Hesselbarth (Quakenbriick, West Germany ). 

Because Alberti’s original description of latedisjunctus is not in- 
formative enough to give reliable diagnostic features, and is not il- 
lustrated, I give here a detailed description of this taxon, based on 
specimens from the type locality, with complete synonymy and addi- 
tional information regarding the type locality. This is a part of my 
Rhopalocera Caucasica Programme having as its aim the compilation in 
one source of a comprehesive and detailed analysis of the recent state, 
origins and zoogeographic features of the Caucasus Region butterfly 

Agriades pyrenaicus latedisjunctus Alberti (1973) 
(Figs. 12-15) 

Lycaena orbitulus Prun. var. dardanus Frr.: Romanoff, 1884, p. 51. 

. pyrenaica var. dardanus Frr.: Bramson, 1890, p. 51. 

orbitulus var. dardanus Frr.: Radde, 1899, p. 420. 

. pyrenaica B.: Egorov, 1903, p. 13. 

orbitulus Prun. var dardanus Frr.: Shaposhnikov, 1904, p. 206. 

. orbitulus Prun. var. dardanus: Alpheraky, 1907, p. 204. 

. orbitulus dardanus (?) Frr.: Riabov, 1926, p. 294. 

. orbitulus var. dardanus Frr.: Warnecke, 1943, p. 175. 

. orbitulus Prun. var. dardanus Frr.: Wojtusiak & Niesiolowski, 1947, p. 58. 
. orbitulus Prun.: Miljanowski, 1964, p. 114. 

. pyrenaica ssp. dardanus: Alberti, 1970, p. 123. 

Polyommatus (Agriades) glandon dardanus Frr.: Korshunoy, 1972, p. 363. 
Lycaena pyrenaica latedisjuncta Alberti: 1973, p. 221. 

eT loti fel fete =! 

General. Upperside wing color closely similar to A. pyrenaicus pyrenaicus, 
differing from dardanus by the more vivid, silvery blue male coloration; dark veins 
are clearly visible on the ground color. Differs from pyrenaicus and dardanus by 
the significant reduction of submarginal spots on the forewing underside, especially 
in males. Female’s wing upperside more abundantly powdered with bright blue 
scales than in both pyrenaicus and dardanus. This character transitional to females 
of pyrenaicus asturiensis Oberthir. Subspecies differs from all other pyrenaicus ssp. 
by male genitalia characters (see text below and Fig. 6). 

Male. Length of forewing (base to tip) 10.0 to 12.5 mm. Upperside of both 
wings of vivid silvery blue shining color, becoming darker toward the margins. This 
darker zone occupies about ¥% of the wing length, and between veins bears diffused 
patches of the ground color. Hindwing bears on its upperside 2 to 3 well developed 
diffused submarginal spots, always present in cells M:-Cu: and M2-Ms, in some 


Figs. 12-15. Agriades pyrenaicus latedisjunctus: 12, 13, ¢ upper and undersides, 
C. Caucasus, Kazbek Mt., 2900-3000 m, 26 July 1972, Y. Nekrutenko; 14, 15, 9 
upper and undersides, same label data. 

specimens also in Cu:-Cuz. Discal spot on the upper side of the forewing always 
contrasting, rounded with a white ring (weakly visible on black-and-white photo- 
graphs). Fringe white, with black strokes at the end of each vein, that do not 
reach the outer margin. Ground color of the forewing underside bright, whitish grey, 
not brownish, somewhat darker toward the base and anal margin. The central 
cell spot in all specimens examined varies from a thin, but contrasting patch to the 
size of a discal spot. Postdiscal spots complete, forming S-shaped row, each spot 
being rounded with a white ring. Submarginal spots incomplete, toward the 
apical part of the forewing gradually disappearing, always present only in cells 
M:-Cux, Cui-Cuz and Cus-2A. A very narrow, precise dark line goes along the 
outer wing margin. Underside of hindwing bears three distinct color zones: distal, 
formed with confluent white postdiscal spots; medial, bright, whitish-grey; and basal, 
bluish grey, with metallic tint. Black markings widely ringed with white, present 
in cells Sc+Ri-Rs (2 spots), Rs-M:, M:-Cu: and Mo-Ms. Discal spot always 
without black pupil. Yellow submarginal lunule in Ms-Cu; shaded with black from 
basal side only; toward the margin gradually transitional into the ground color, some 
specimens bear a black pupil at this point. 

Female. Length of forewing (base to tip) ranged from 10.0 to 12.5 mm. 
Ground color of the upperside of both wings dark, brown-grey. Forewing bears 
discal spot of deep black color, ringed with white broad circle, with characteristic 
drawing off toward the outer margin. Discal spots on the hindwing upperside 
variable: from almost complete disappearance to the size of the forewing discal 
spot. Hindwings bear on their upperside diffused submarginal spots as in males. 
Wings of many females bear bright diffused postdiscal and submarginal spots of 
the male color, often with greenish tint. Underside color and pattern as in male, 
but ground color more vivid, brownish, markings developed more strongly. 

VoLUME 28, NUMBER 3 285 

Figs. 16-19. Agriades pyrenaicus latedisjunctus, genitalia: 16, male, general view, 
aedeagus removed; 17, 18, male, aedeagus, lateral and dorsal projections; 19, female, 
general view, ventral projection. 

Male genitalia (Figs. 6, 16-18). General appearance as in all other Agriades. 
Juxta horseshoe-shaped, with divergent upper extremities, strongly chitinized. The tip 
of the upper valval lobe obtuse, oblique (in dardanus rounded, symmetric—see Fig. 
5), bears 15 to 21 teeth (20 specimens dissected). The isthmus between the body 
of valva and the tip broad, poorly expressed. 

Female genitalia (Fig. 19). I have found no feature of diagnostic value in 
the female genitalic armatures in all specimens of all species of Agriades ever 
examined. The female genitalia of latedisjunctus are figured here to complete the 
description and this figure covers all Agriades. 

Material studied. 49 ¢¢, 10292, C. Caucasus, Georgian Soviet Socialist 
Republic, Kazbek Mt., 2900-3000 m, 26 July 1972 (Y. Nekrutenko); 12 6 6,3 2 @Q, 
Kazbegi circ., 1850 m, 24 July 1972 (Y. Nekrutenko); 3 ¢ 6, Abkhasia, Mzy (Mzym) 
Lake, 2300 m, 12 July 1972 (Y. Nekrutenko); 2 66, 1 9, Abkhasia, Awadhara, 
2000-2200 m, July-August 1967 (E. Miljanowski); 1 ¢, Georgia, Lebarde, 8 
June 1962, E. Didmanidze (coll. S. Miljanowski); 2 ¢ 6, Teberda, N. W. Caucasus, 
July 1935, L. Sheljuzhko (Zool. Mus. Kiev Univ.); 4 66, 3 29, Daghestan, 


Fig. 20. Agriades pyrenaicus latedisjunctus, type locality. Upper alpine zone on 
the Eastern slope of Kazbek Mt. at an elevation of 2900-3000 m. 

Levashi, June 1926, M. Riabov (Zool. Mus. Kiev Univ.); 2 ¢ ¢, Tskhra-Tskaro, 
Borzhomi, Caucasus Minor, 2520 m, July 1914, L. Sheljuzhko (Zool. Mus. Kiev 
Univ.); 1 ¢, Armenia, Amamly (subalpine zone), 20 July 1925, M. Riabov (Zool. 
Mus. Kiev Univ.); 1 ¢, Armenia, Alagéz Mt., 15 May (?) 1935, B. Tkatshukov 
(Zool. Mus. Kiev Univ.). 

Type locality (Fig. 20). In addition to the data given by Alberti (1973), the 
type locality should be restricted to the area on the Eastern slope of the Kazbek 
Mt., where the butterflies are most abundant. This place is situated between the 
Tsminda Sameba (St. Trinity) church over the Gergeti village and the Gergetskiy 
glacier tongue margin and fore moraine. It is in an hour or two of rather easy 
climbing from the Georgian Military Highway at Gergeti village, on the left bank 
of Terek river. In the Kazbegi village vicinity on the opposite side of Terek (1850— 
1900 m) the butterfly is rather scarce. 


I wish to acknowledge the generous help I received from Mr. T. G. 
Howarth of the British Museum (Natural History), and from Dr. W. 
Forster of the Zoologische Sammlung des Bayerischen Staates in Munich, 
who supplied me with many literary sources from their libraries. Mr. 
Gerhard Hesselbarth collected and kindly forwarded material, including 
the type series, of the new subspecies named here in his honor; compara- 
tive materials have been obtained from Dr. F. Fernandez-Rubio, Dr. 

VoLUME 28, NUMBER 3 287 

Hans Malicky and Mr. Colin W. Wyatt. I am grateful to Dr. Tamara 
Zrazhevska for the use of her microscopical facilities. Drafts of this 
paper have been discussed with Mr. Yuri P. Korshunov (Novosibirsk ) 
and Dr. Eugene S. Miljanowski (Sukhumi). My thanks are due to Dr. 
Theodore D. Sargent who corrected and edited the manuscript. 


ALBERTI, B. 1970. Vergleichende Eindriicke von der Lepidopterenfauna des Nord- 
und Siidkaukasus sowie Transkaukasien. Nachr. Bayer. Entomol. 19 (6): 118- 

1973. Ergiinzende Bemerkungen zu Higgins & Riley: “A field guide to 
the butterflies of Britain and Europe,” nebst Beschreibung der Lycaena 
pyrenaica latedisjuncta n. subsp. Entomol. Zeitschr. 83 (19): 217-223. 

BETHUNE-BAKER, G. T. 1913. Notes on the specific distinction of certain species 
in the orbitulus and pheretiades section of the genus Plebeius. Trans. Entomol. 
Soc. Lond.: 205-212, pls. VI-VIII. 

Bramson, K. L. 1890. Die Tagfalter (Rhopalocera) Europas und des Caucasus, 
analytisch bearbeitet von K. L. Bramson. Verlag des Verfassers, Kiew. 150 p., 

Buresu, I. & K. Tutesuxoyv. 1930. Horisontal distribution of the Lepidoptera in 
Bulgaria. Izvestia na Tsentralnite Prirodonauchni Instituti 2: 124-125 (In 
Bulgarian ). 

CHAPMAN, T. A. 1908. Erebia lefebvrei and Lycaena pyrenaica. Trans. Entomol. 
Soc. Lond.: 314-316, pls. XI—XIII. 

Ecoroy, N. 1903. Lepidoptera of the Northern slope of the Central Caucasus. 
Izvestia Kavkazskogo Otdela Imperatorskogo Russkogo Geograficheskogo 
Obshchestva 16 (1): 9-24 (In Russian). 

FERNANDEZ-Rupio, F. 1972. Redescubrimiento de una rara mariposa en Sierra 
Nevada. Note sobre la captura del Lycaenido: Plebejus gladon ziillichi 
Hemming, 1933 (= nevadensis TZiillich y Reisser, 1928). Arch. Inst. 
Aclimatacién 15: 161-167, lams. II-V. Almeria. 

Forster, W. 1938. Das System der palaarktischen Polyommatini (Lep., Lycaen.). 
Mitt. Miinchn. Entomol. Ges. 28(2): 97-118. 

FREYER, C. F. 1845. Neuere Beitrége zur Schmetterlingskunde mit Abbildungen 
nach der Natur 5: 59-60, Tab. 419, figs. 2-3. Augsburg. 

GERHARD, B. 1853. Versuch einer Monographie der Lycaenen als Beitrag zur 
Schmetterlingskunde mit Abbildungen nach der Natur 10: 11, pl. 17-18. 

Hremminec, F. 1967. The generic names of the butterflies and their type species. 
Bull. Br. Mus. Nat. Hist. (Entomol.) Suppl. 9: 31-32. 

Hiceins, L. G. & N. D. Rumsey. 1970. A Field guide to the Butterflies of Britain 
and Europe: 286-288. Collins, London. 

Korsuunov, Y. P. 1972. A catalogue of the Lepidoptera Rhopalocera of the fauna 
of U.S.S.R. II. Entomologicheskoye Obozreniye 51(2): 352-368 (In Russian). 

Kuznecov, N. J. 1929. Insectes Lépidoptéres. Faune de VURSS et des pays 
limitrophes 1(2): DLXXIII. Leningrad. (In Russian). 

Lattin, G. De. 1950. Tirkische Lepidopteren I. Rev. Facult. Sci. Univ. Istanbul, 
ser. B 15(4): 301-331. 

MityAnowski, E. S. 1964. The butterfly and moth fauna of Abkhasia. Trudy 
Sukhumskoi Opytnoi Stantsii Efirnomaslichnykh Kultur 5: 91-190 (In Russian). 

Miter, L. D. 1969. Nomenclature of wing veins and cells. J. Res. Lepid. 8(2): 


NEKRUTENKO, Y. P. 1972. A new subspecies of Eumedonia eumedon (Lycaenidae) 
from Caucasus. J. Lepid. Soc. 26(4): 210-212. 

OseRTHUR, C. 1910. Notes pour servir a établir la faune Francaise et Algérienne 
des Lépidoptéres. Suite. Etudes de lépidoptérologie comparée 5: 1-641. 

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PAU 6: 1-74. 


Plant resistance to insect attack has been studied largely in connection with 
agricultural practices and crop plant breeding (Beck 1965, Ann. Rev. Entomol. 
10: 207-232), although the principles gained therefrom should apply to natural 
situations as well. Butterfly larval foodplants in the wild likewise have probably 
developed strains that are resistant to attack. This fact would account for spotty or 
discontinuous distributions of some species, although the effect would be difficult 
to distinguish from extinction due to other causes. In the field, one frequently en- 
counters areas where a known foodplant is present but the butterfly is absent. E.g., 
Papilio indra fordi Comstock & Martin feeds on Cymopterus panamintensis Coult. & 
Rose but not on the subspecies acutifolius (Coult. & Rose) Munz (Shields, Emmel, 
& Breedlove 1969, J. Res. Lepid. 8: 21-36). Toxic secondary plant substances 
may act as repellents; ecdysone or juvenile hormone or their analogues in plants may 
protect them from attack (Fraenkel 1969, Entomol. Exp. Appl. 12: 473-486; Hsiao 
1969, Entomol. Exp. Appl. 12: 777-788). Plant resistance can disturb the insect’s 
normal behavior, growth, and survival (Beck, 1965). 

OAKLEY SHIELDS, Department of Entomology, University of California, Davis, 
California 95616. 

VoLUME 28, NUMBER 3 289 


John H. Masters’ very thought-provoking and controversial paper under the above 
title (1972, J. Lepid. Soc. 26: 249-260) cannot be allowed to pass without comment. 
In the first place I consider it utterly wrong for any section of entomologists, be they 
lepidopterists, coleopterists, dipterists or any other, to attempt to formulate a code 
that would apply to their own Order only. Any such code must apply to all 
Orders of insects. Nor do I think it right that the requirements of the geneticists 
should be dismissed in such a cavalier fashion. 

It would, perhaps, be most convenient if I listed my comments under the same 
headings as used in the original article. 

INFRASUBSPECIFIC VARIATION (p. 250). Masters writes, “Other than a 
general agreement that infrasubspecific names should not be placed in italics. . . .” 
But is this true? It certainly is not for the four British entomological journals to 
which I subscribe, and the British Museum (Natural History) continues to print 
infrasubspecific names in italics in its Bulletin (Entomology). Again, is it true 
to say, “there has been a very sharp decline in the publication of formal names 
to apply to infrasubspecific varieties in the last twenty years,’ and, “most authors 
are content to describe examples of infrasubspecific variations without attempting 
formally to name them”? The first may be partially true, probably because most of 
the well marked variations have already been described and named, but in my 
opinion, the second is not, and, in any case, what is the point of a description without 
attaching a name to it? Which is the more preferable title for a hypothetical 
article, “The genetics of Arctia caja L. and its form. .. . or “The genetics of 
Arctia caja L. and its form as described in 1970, Entomologist, ...:... I know 
which I would prefer, and I think the majority of entomologists would agree with 
me. I have covered the question of Lepidopterists ‘going it alone’ in my introductory 

definition of polymorphism is undoubtedly scientifically correct, it does appear to 
reduce the proportion of the rarer to the commoner form to far below what is 
normally considered as polymorphism. Surely there must be a point, well illustrated 
by Industrial Melanism in Britain, when a form ceases to be a mere mutant and 
becomes polymorphic. To take the geometer Biston betularia L. and its black form 
carbonaria Dbl. as an example, in the late eighteen hundreds and early in the 
twentieth century the black form was a great rarity, possibly so rare that it could 
not be maintained except by recurrent mutation, chiefly because its colour made 
it overconspicuous when at rest and it suffered heavily from predators. Once 
industrial pollution had altered the environment, the position was reversed and it 
was the typical speckled form that was at a disadvantage and, as a result, the 
black form, which was genetically dominant, rapidly increased its proportion of the 
total population until it is the prevalent form in many areas today. 

Whilst there is some point in applying a nomen collectivum to all the forms in 
a group that are a manifestation of the same gene, it must not be forgotten that 
what may appear to be similar forms, even in the same species, may be the 
result of completely different genes. Whilst accepting the nomen collectivum in 
limited cases, I think there is still a need for a formal name for the various forms, 
and I also consider that the addition of the author’s name is essential, not, as Mr. 
Masters points out, as a compliment to the author but to pinpoint the reference. 

The suggestion of applying the model’s name prefixed by pseudo- to the various 
forms of polymorphic mimics is only a partial solution of the problem. How, for 
example, are the four forms of Danaus chrysippus L., viz. chrysippus L., alcippus 
Cr., dorippus Klug and albinus Lanz, to be treated and what about the many 
examples of polymorphism in procryptic moths, such as Achaea lienardi Bsd., A. 


praestans Mab., Blenina quadripuncta Hamps. and Odontodes aleuca Guen., to name 
only a few. Here, again, formal names seem to be the only answer. 

Many aberrations in the genus Parnassius, the Lycaenidae and Arctiidae are almost 
certainly multifactorial in origin and, overlapping as they do, are probably best 
treated with descriptive, as opposed to formal, names. The use of descriptive terms 
for aberrations was probably carried to the extreme in Bright & Leeds Monograph 
of the British Aberrations of the Chalk Hill Blue Butterfly, Lysandra coridon (Poda) 
1761. (Bournemouth 1938) which described some four hundred types of aberration. 

MUTANT OR ABERRATIONAL FORMS (p. 253-254). The reference to the 
effect of cold on the pupae of Euphydryas phaeton (Drury) raises an interesting 
point. Normally the effect of unusual temperatures is an interference with the 
normal process of pigmentation, and Haggett (1952, Entomologist) has shewn 
that a number of the named forms of Rhodometra sacraria L. are the result of low 
temperatures on the pupa, examples carrying the factor for redness producing f. 
sanguinaria Esper at slightly lower temperatures and f. rosea Oberthur at the lowest 
possible, whilst those without the factor for redness produce f. labda Cr. at slightly 
lower temperatures and f. atrifasciaria Stephens at the lowest possible. In other 
words the visible effect of the gene is enhanced by low temperatures, probably 
through the greater length of the pupal period. In the arctiid Panaxia dominula L., 
it has been established that there are certain genes that do not manifest them- 
selves unless the pupa is exposed to abnormally low temperatures. Whilst I agree 
that purely temperature forms are not worthy of a name, I do think there is 
a case for naming forms which are a combination of temperature and a specific gene. 

I cannot agree with Masters’ statement, “Whether genetic or non-genetic in cause, 
aberrants are not normally an integral part of any population, each specimen is an 
individual without direct connection with any succeeding individual that may 
resemble it.” This is manifestly not correct in the case of genetical aberrants, even 
if the gene is fully dominant and lethal when homozygous it will survive unless the 
heterozygotes are at such a disadvantage that all are killed by predators, and a rare 
and recessive gene can survive undetected for generations in heterozygotes. An 
illustration of this occurred here recently, three specimens of an aberration of 
Charaxes brutus Cr., lacking the chestnut component of the underside basal markings, 
were trapped in the same area and within a few days of each other and were 
fairly obviously the progeny of one female. It is only a matter of time before a 
pairing between two apparently normal individuals, but both heterozygous for 
this particular gene, occurs and the aberration re-appears. 

I agree that gynandromorphs, somatic mosaics and other freaks are best left 
unnamed, but it must not be forgotten that many of these are genetic in origin. 

SEASONAL FORMS (p. 254-255). Here is one of the few parts of the paper 
with which I am in partial agreement. I say ‘partial’ as I do not care for Masters’ 
third, and preferred, alternative. I feel that the second is by far the best. Numerals 
or letters to denote seasonal forms rather break down when applied to wet and dry 
forms in the tropics, a wet form may occur earlier in the year in one part of a 
species’ range and later in another. 

HYBRIDS (p. 255-256). Here again I am only in partial agreement. Whilst 
accepting the first three classes and the method of naming them, I feel that once 
a stable hybrid population has established itself in nature it is far preferable to give 
it a name and treat it as a species, for that is undoubtedly what it will become, 
if not sooner then later. Papilio kahli Chermock & Chermock may be a fairly straight- 
forward case, but Warren’s hybrid Pieris, based mainly on deformed andraconia, 
is very much a matter of opinion and is unlikely, in my opinion, ever to be proved 
conclusively. After all, many so-called subspecies are probably nothing more than 
hybrids between two separate subspecies that have met and then become isolated. 

D. G. SEvASTOPULO, F.R.E.S., P.O. Box 95026, Mombasa, Kenya. 

VoLUME 28, NUMBER 3 291 


Recent reports of an attempted interfamilial mating (Shapiro 1973, J. Lepid. Soc. 
27: 159) and an interfamilial courtship (Shapiro 1972, J. Res. Lepid. 11: 197-198) 
suggest these may occur at least as frequently as the rare pairings and courtships 
of sympatric congeners, which have received some attention in the literature (Downey 
1962, J. Lepid. Soc. 16: 235-237). Another recent study of butterfly mating 
behavior (Scott 1972 [1973], J. Res. Lepid. 11: 99-127) has provided welcome 
data for verifying such attempts at copulation by behavioral traits. The purpose 
of this note is to report another attempted interfamilial mating—¢ Lycaena phlaeas 
americana Harris (Lycaenidae) and @ Phyciodes tharos tharos (Drury) (Nymphali- 
dae )—and to comment on its significance. 

The instance occurred between 1522 and 1528 hrs. on a lawn in New Paltz 
(Ulster County), New York, on 17 September 1973. While observing mating behavior 
of a number of P. tharos at this site, I particularly noticed one pair attempting 
copulation atop a clump of grass. It was a fresh female P. tharos and fresh male 
LL. phlaeas. The latter was approximately 8 mm smaller in expanse than the former. 
The male, in the characteristic position behind and facing the same direction as the 
female, made repeated attempts at genital contact by arching its abdomen beneath 
and to (what appeared to be) both sides. The female remained docile, wings 
horizontal except for a slow, occasional fanning to an angle of about 30 or 40 
degrees. The male held its wings at a 45 degree angle throughout. Having no 
success at contact, the male moved forward until its head and forelegs were atop the 
female’s abdomen. This apparently startled the female, which flew lazily away 
to a site about 1.5 m away. The male followed, slowly, and similar behavior ensued 
at the second site. For an unapparent reason the female then flew to a third site, 
very near the first. The male followed, but this time became quite pugnacious and 
upon aggressively approaching the female caused her to fly off quickly. The male 
was unable to follow and was collected for sexual verification. 

These species are phenotypically similar: both exhibit predominantly orange and 
black wing characters, mostly in “spotted” patterns, and males of both species are 
usually smaller than the female, as in the case of this attempted pairing. Further, 
both species seek mates by “patrolling” (Scott, loc. cit.). The female P. tharos 
was apparently receptive, displaying none of the rejection postures known to butter- 
flies, but exhibiting instead the stationary and “basking” behaviors often mentioned 
as receptive traits (Scott, loc. cit.). She flew off only after notable pugnacity on 
the part of the male, a fact which may be doubly significant since both species 
are noted for this aggressive trait (Klots 1951, A Field Guide To The Butterflies, 
Houghton Mifflin, Boston). 

Females of Lycaena helloides (Boisduval) reportedly fan their wings as a 
receptive trait (Shapiro 1973, loc. cit.), and if L. phlaeas females do likewise, this 
might have encouraged the male L. phlaeas’ advances. Two attempted matings 
of L. phlaeas were noted at the same locality at 1540 and 1600 hrs. Characteristic of 
these was apparent rejection behavior by the female (wings closed tightly above the 
thorax, and a quick “waddling” through the grass) and extreme pugnacity by the 
male (following quickly behind, trying to “steer” the female into an appropriate 
mating position). All of these observations support the conclusion that the P. 
tharos female and L. phlaeas male noted above were attempting copulation. 

Scott (loc. cit.) states that coloration, movement, and size are important to the 
visual components of butterfly mating. Shapiro (1972, 1973, loc. cit.) notes the 
evident importance of phenotypic similarities (and also pheromones) in eliciting 
such mating mistakes. He discusses the surprising phenotypic dissimilarities of his 
interfamilial “mates.” As with his species, the pheromones of P. tharos and L. 
phlaeas have not been studied. If eventual pheromone data do not indicate other- 


wise, this attempted mating of L. phlaeas and P. tharos may represent a more 
“classic” example of similar phenotypes eliciting an attempted interfamilial mating— 
the type which would seem most probable if such events do occur more frequently 
than lepidopterists have suspected. 

I would like to thank Br. (Dr.) Adam McCoy, Holy Cross, for editorial assistance. 

Kurt JoHunson, (Br.) Novitiate, Order of the Holy Cross, West Park, New York 
12493, and Museum Research Associate, Museum of Natural History, University 
of Wisconsin, Stevens Point 54481. 


Glyphipterygid moths are diurnal and usually associated with blooming plants 
favored by the particular species, in addition to their hostplant. Reports of 
glyphipterygids at lights are as infrequent as for other diurnal insects and only 
Tortyra slossonia (Fernald), Choreutis carduiella Kearfott, and a Glyphipteryx sp. 
have been sparingly encountered this way, in addition to what is tentatively identified 
as Choreutis leucobasis Fernald. These Florida reports, however, involve only one 
or two individuals at a time, as do light collection records of Anthophila pariana 
(Clerck) from the Northeast. The T. slossonia records are mainly from light trap 
collections made by Mrs. Spencer Kemp on Key Largo and also involve only one 
or two specimens some nights. 

Collections of diurnal insects at light have been attributed to the fact that the 
light has been set up near the resting place of the insect which moves to the 
light upon being disturbed. The large number (70+) of Tortyra slossonia collected 
at a blacklight near Tavernier, Key Largo, the evening of 20 June 1973 from 
about 2000 to 2300 hours indicates that it may be nocturnally active unlike other 
glyphipterygids. Two nights earlier on the north end of Key Largo, about 12 T. 
slosscnia moths were also taken at a blacklight. 

(Florida Agricultural Experiment Station Journal Series No. 5275.) 

Joun B. HEpPPNER, Department of Entomology and Nematology, University of 
Florida, Gainesville, Florida 32611. 


A worn male specimen of the neotropical day-flying moth, Urania fulgens Walk. 
(Uranidae), was captured by V. J. Farkas in downtown Fort Walton Beach, along 
Santa Rosa Sound, on the Gulf of Mexico side of northern Florida, at 1400 hrs. 
on 9 September 1973. It was hovering over a lantana bush in a weedy summer- 
cottage area. A common migratory species in Yucatan and mainland Mexico, this 
specimen was probably blown northeast to Florida by tropical storm “Delia” which 
passed over the Yucatan Peninsula around 5 September and then continued into 
the Gulf. This appears to be a new record for Florida (not listed in Kimball, 1965, 
Lepidoptera of Florida, Florida Department of Agriculture) and for the eastern 
United States. 

THomas C. EMMEL, Department of Zoology, University of Florida, Gainesville, 
Florida 32601. 
V. J. Farkas, 722 Hollywood Boulevard, Mary Esther, Florida 32569. 

VOLUME 28, NUMBER 3 293 


Romualdo Ferreira d’Almeida, son of Henrique Ferreira de Almeida 
and Izabel Pereira de Almeida, was born in Rio de Janeiro on 12 February 
1891 and died there on 24 August 1969. He married Aida Moreira dos 
Santos and had four sons, Nelson, Nysio, Newton and Ney. 

His life-long interest in butterflies started at an early age, as did his 
interest in music which he inherited from his father. His first earnings, 
which he soon spent on his collection, were frequently made by playing 
the organ in church. 

Needing to have a reliable source of income, he applied for govern- 
ment service and was accepted as an assistant cleaner to the Director 
General of Post Offices on 15 February 1917. Three years later he was 
made a second class cleaner and on 14 April 1921 he was promoted to 
delivering mail, in a third class capacity; he was promoted to the second 
class on 24 December 1934. 

He thus worked every afternoon in order to have a reliable, if modest, 
source of income, and left the mornings free to dedicate himself to 
his passionate interest. His collection, at first very small due to lack 
of space and working conditions, and done without any outside help 
whatsoever, grew slowly. Because of his lack of support in Brazil, he 


started corresponding with entomologists in other countries, first in 
France and Germany and later throughout the world. In Brazil he 
remainded unrecognized and rejected by all the research institutions 
which he contacted, until a friend of his, Sr. J. Pinto, a photographer 
at the Oswaldo Cruz Institute in Rio de Janeiro, introduced him to Dr. 
Lauro Pereira Travassos, also of the Oswaldo Cruz Institute, in 1933. 
Despite having 24 publications in French and German, Romualdo F. 
d’Almeida had been ignored for 20 years. Dr. Travassos, a specialist in 
helminths and Lepidoptera, soon realized that he was dealing with 
someone worthy of recognition, and thus, with the support of the in- 
fluential deputy, Arthur Neiva, arranged for Romualdo F. d’Almeida to 
deliver the mail within the Oswaldo Cruz Institue. Here, with good 
working conditions, equipment, a specialized library, and a suitable 
atmosphere to work in, he was able to improve and increase his output. 
He gave up delivering mail. Here he wrote his largest and best-known 
works, the revisions of the genus Eurema ( Pieridae), the genus Actinote 
(Nymphalidae) in the southeast of Brazil, and on the family Danaidae; 
he also worked out his plans for the research that he was to do throughout 
his life. From this time on, Romualdo F. d’Almeida was recognized and 
respected in Brazil. 

In 1937, he had the opportunity of accompanying the border Com- 
mission (northern sector), led by Commander Braz Dias de Aguiar, 
when an outstanding collection of butterflies was made in the area of the 
Cumina and Trombetas rivers in Para. Another important expedition, 
made only shortly before he died, was to the Amapa Territory in 1967; 
this was at his own expense and he spent all of the small amount of money 
which he possessed. 

He remained at the Oswaldo Cruz Institute until 1 December 1940, 
officially delivering letters but in practice studying butterflies. From 
2 December on, at the invitation of Dr. Salvador de Toledo Piza, he was 
appointed as an assistant in the Zoology Department of the Secretary of 
Agriculture, Industry and Commerce for the State of SAo Paulo, and was 
thus under the directorship of Dr. Oliverio Pinto. This was the first time 
that he had been employed as a research worker. Here, not having a 
collection, with which he would have been much happier, his research 
was mainly bibliographical, including various aspects of nomenclature. 
He remained there until 4 July 1944, when he was transferred to an 
appointment as Assistant Naturalist at the Ministry of Education and 
Health, at the initiative of the director of the National Museum of Rio 
de Janeiro, Dr. Heloisa Alberto Torres. 

On his return to Rio, he was able to go back to working on his own 

VOLUME 28, NUMBER 3 295 

collection. Now, with a responsible job and a good salary, he was un- 
fortunately at the end of his career, being already 53 years old; however 
he worked unceasingly until] his retirement in 1967, and after this worked 
at home until the time of his death. 

Without doubt Romualdo F. d’Almeida knew the Lepidoptera of South 
America better than anyone else. His scientific output was considerable 
and it is not known why his first publication in 1913 was only discovered 
in Brazil 20 years after it had been written. All this time he had been 
unknown, and had been forced to study Latin, French and 
German in order to write and publish his work. His works 
covered all groups of the Rhopalocera and some few Heteroc- 
era, mainly concerning nomenclature, systematics and_ biology. 
The group he was most interested in was the Ithomiidae but he never 
accomplished his ambition to revise this family. In spite of having had 
to struggle against so many difficulties, having had a large family, and 
having received little moral or material support, he was perhaps the 
most productive Brazilian entomologist. He left a collection of more 
than 27,000 specimens in excellent condition, almost all of which had 
been identified; he wrote 112 scientific publications, a list of which, with 
comments, will be published by F. Martin Brown in an early issue of this 

The author had the pleasure of being a student of Romualdo d’Almeida 
between 1957 and 1969, during which period he visited Romualdo 
d Almeida at home frequently, to learn about the morphology, systematics 
and biology of Lepidoptera. At this time, because of his poor health, he 
seldom left his house to go to the National Museum in Rio, and when 
he did, it was only to consult various books or to visit his friends. Even 
so, he would go to the field to collect something which might be of 
interest to his work and always used to be pleased when he found any 
species which he did not have in his collection, although this was a rare 
occurrence. He had a persistent character and did not like to ask favours 
from friends; he also hated bureaucracy and was in the habit of using 
the backs of official forms for writing his scientific papers. He used to 
work on his collection daily in his house and died putting the finishing 
touches to his bibliographic catalogue of the Ithomiidae. 

He was a member of various scientific societies: The Lepidopterists’ 
Society (of which he was vice president in 1952), corresponding member 
of the Sociedade entomologica Argentina; corresponding Academician of 
the Academia Chilena de Ciencias Naturales; corresponding member of 
Sociedade Entomologica de Chile; member of the Societe entomologique 
de France, from which he received the Alcides d’Orbigny prize in 1929; 


the Societé Linean de Lyon; L’'Union de Entomologiste Belges; Inter- 
nationaler Entomologischer Verein, Frankfurt; and the Sociedade 
Brasileira de Entomologia, which dedicated a book to him in 1945, In 
1950, Romualdo Ferreira d’Almeida was awarded the medal of the 
“Oficial da Ordem Nacional do Merito” by Getulio Vargas, the President 
of Brazil, and later received the medal from President Eurico Gaspar 

I am grateful to Dr. Judith Smith of Universidade Federal do Parana, 
who kindly translated the text from Portuguese to English. 

Pror. Ouar Micke, Univ. Fed. Parana, Curitiba, Parana, Brazil. 


A female specimen of the Tailed Sulphur, Eurema proterpia Fabr., was taken 
seated on wild aster blossoms in a field near Rantoul Gap, nine miles east of Ottawa 
in Franklin County, Kansas, on 15 October 1973. I netted the specimen just a few 
feet away from a specimen of the Mexican Snout Butterfly, Libytheana carinenta 
(Cramer) which was also seated on the asters. Neither butterfly has ever been 
recorded in Kansas before and I presume both of them to be new state records 
(although carinenta was taken by the dozens in this area during the autumn of 
1971). Both specimens were somewhat worn and are presumed to have been 
migrants entering the region from farther south. Both specimens will be deposited 
in the Los Angeles County Museum at Exposition Park, Los Angeles, California. 

WitiiaAM H. Howe, 822 East Eleventh Street, Ottawa, Kansas 66067. 

Editor: THEODORE D. SARGENT, Department of Zoology, 
University of Massachusetts, Amherst, Massachusetts 01002 

K. S. Brown, J. M. Burns, R. H. Carcasson, J. P. DONAHUE, 
J. F. Gates Criarxe, C. D. Ferris, R. O. KENDALL, H. K. CLENcH, 
J. H. Masters, L. D. Mruter, A. P. Pratt, A. M. SHapio, J. R. G. 


Contributions to the Journal may deal with any aspect of the collection and study 
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Text: Manuscripts should be submitted in duplicate, and must be typewritten, 
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Literature Cited: References in the text of articles should be given as, Sheppard 
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under the heading Lirerature CirTep, in the following format: 

SHEPPARD, P. M. 1959. Natural Selection and Heredity. 2nd. ed. Hutchinson, 
| London. 209 p. 
196la. Some contributions to population genetics resulting from the 
study of the Lepidoptera. Adv. Genet. 10: 165-216. 
In the case of general notes, references should be given in the text as, Sheppard 
(1961, Adv. Genet. 10: 165-216) or (Sheppard 1961, Sym. Roy. Entomol. Soc. 
London 1: 23-30). 

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(Continued from outside front cover) 

pAE).. Bryant Mather. 2 220 
Ex Satvapor (NyYMPHALIDAE). Alberto Muyshondt and 
Alberto Muyshondt, Jr, 0) 224 
Clifford. D.. Ferris, 20 230 

TEXAS AND THE Unirep States. Roy O. Kendall ______.___ 243 
Knudson, 800 0 ee 246 
O. Kendall. a ee 249 
Charles V. Covell, Jr 2 253 
A Costa Rican Mountain Forest. Allen M. Young ___ 257 
Pest Crossrs THE ATLANTIC (PrermDAe). Brian O. C. Gardiner 269 

Technique for specific determinations of dead pupae of Eupithecia 
(Geometridae), K. B. Bolte 0200000000000 204 
New state records for Indiana and Illinois. Irwin Leeuw ..........2-....-.-------- 211 
Phyciodes mylitta (Nymphalidae) on Vancouver Island. Richard Guppy 223 
“Attacks” by Polygonia interrogationis (Nymphalidae) on chimney swifts 

and insects. D. Paul Hendricks | _...:..00...002.0..2.\))) 9 236 
Phyciodes texana (Nymphalidae) in California. Richard C. Priestaf ...........- 242, 
Butterflies attracted to amber glass. Richard Guppy ......--..-.-..:--:.-000e-000--= 248 - 
Pellicia costimacula Herrich-Schaffer in the United States (Hesperiidae). 

Mike A. Rickard (occ 252 
Celastrina ebenina (Lycaenidae) in North Carolina. Richard E. Price, Jr. 268 
Resistance in butterfly foodplants. Oakley Shields .......2....220..121:120eee-00--0e 288 
A proposal for the uniform treatment of infrasubspecific variation we 

lepidopterists. D. G. Sevastopulo ............00........... 289 
An attempted interfamilial mating (Lycaenidae—Nymphalidae). Kurt 

FORNSOR ee 291 

Tortyra slossonia collected at UV light on Key Largo, Florida (Glyphi- 
pterygidae). John B. Heppner. 2....0....0.222..-s0sslse re 
Urania fulgens (Uranidae) captured in Florida. Thomas C. Emmel and 
Weeds Parkas ni ee 
Eurema proterpia (Pieridae) in Kansas. William H. Howe ...........22------------- 

OBITUARY, (28 Oe Oe Ted) a ea 

Volume 28 1974 Number 4 


of the 


Published quarterly by THE LEPIDOPTERISTS’ SOCIETY 

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ie 27 December 1974 


Harry K. CLeNcH (Pittsburgh, Penn.) President 

ANDRE BLANCHARD (Houston, Texas) President-elect 
RonaLp W. Honces (Washington, D.C.) Ist Vice President 
J. C. E. Riorre (Toronto, Ontario) Vice President 

L. Vari (Pretoria, South Africa) Vice President 

S. S. NicoLay (Virginia Beach, Va.) Treasurer 

Lee D. Mitter (Sarasota, Florida) Secretary 

Members at large (three year term): R. O. Kenpatx (San Antonio, Tex.) 1975 
J. M. Burns (Cambridge, Mass.) 1974 J. A. Powetu (Berkeley, Calif.) 1975 
R. H. Carcasson (Vancouver, B.C.) 1974 J. T. Brewer (Auburndale, Mass.) 1976 
M. C. Nrevsen (Lansing, Mich.) 1974 K. S. Brown (Rio de Janeiro, Brazil) 1976 
D. C. FErcuson (Washington, D.C.) 1975 K. W. Pump (Fairbanks, Alaska) 1976 

The object of the Lepidopterists’ Society, which was formed in May, 1947 and 
formally constituted in December, 1950, is “to promote the science of lepidopterology 
in all its branches, .... to issue a periodical and other publications on Lepidoptera, 
to facilitate the exchange of specimens and ideas by both the professional worker and 
the amateur in the field; to secure cooperation in all measures” directed towards 
these aims. 

Membership in the Society is open to all persons interested in the study of 
Lepidoptera. All members receive the Journal and the News of the Lepidopterists 
Society. Institutions may subscribe to the Journal but may not become members. 
Prospective members should send to the Treasurer full dues for the current year, 
together with their full name, address, and special lepidopterological interests. In 
alternate years a list of members of the Society is issued, with addresses and special 
interests. There are four numbers in each volume of the Journal, scheduled for 
February, May, August and November, and six numbers of the News each year. 

Active members—annual dues $10.00 
Student members—annual dues $7.50 
Sustaining members—annual dues $20.00 
Life members—single sum $150.00 
Institutional subscriptions—annual $15.00 

Send remittances, payable to The Lepidopterists’ Society, and address changes to: 
S. S. Nicolay, 1500 Wakefield Dr., Virginia Beach, Virginia 23455. 

Memoirs of the Lepidopterists’ Society, No. 1 (Feb. 1964) 

by Cyrit F. pos Passos 

Price, postpaid: Society members—$5.00, others—$7.50; uncut, unbound signatures 
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Volume 28 1974 Number 4 



Systematic Entomology Laboratory, Agricultural Research Service, USDA, 
c/o U.S. National Museum, Smithsonian Institution, Washington, D.C. 20560 

Moths identified as Semiothisa (or Philobia) aemulataria (Walker) in 
most collections from the southeastern United States were found to be 
a mixture of two species of extremely similar appearance. One of these 
is the true aemulataria; the other is undescribed. Semiothisa aemulataria 
is common in collections and widely distributed, occurring across southern 
Canada from Newfoundland to Alberta and southward to the Gulf States, 
including northern Florida and eastern Texas. Southern specimens tend 
to be smaller, darker, and less clearly marked than northern ones, but I 
found no structural differences and continue to regard such variants as 
belonging to the same species. 

The undescribed species that has been confused with aemulataria is 
somewhat larger, generally paler than southern examples of aemulataria 
from the same region, and with the markings more boldly defined, 
especially the intense, red-brown postmedial bands on the undersides of 
both wings. The heavily swollen (incrassated) hind tibia of the male 
(Fig. 14) at once distinguishes it from all of the North American species 
of Semiothisa Hubner formerly placed in Philobia Duponchel (aemulataria 
(Walker), and versitata, perplexata, aspirata and ulsterata (Pearsall) ). 
The widely sympatric Semiothisa aequiferaria (Walker), often confused 
with aemulataria in the South, also has a swollen male hind tibia, but 
the moth is smaller and darker, with the outer margin of the forewing 
less obivously notched behind the apex. The new species, which I am 
naming Semiothisa promiscuata, occurs from Maryland to Florida, and 
west to Illinois, Arkansas and eastern Texas. Its hostplant and early 
stages are unknown. | 

Semiothisa aemulataria was described as Macaria aemulataria Walker 
(1861: 884) from one male and one female in the British Museum 


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Figs. 1-6. Specimens: (1) Semiothisa promiscuata, n.sp., holotype; (2) same 
specimen, underside; (3) S. promiscuata, allotype; (4) same specimen, underside; 
(5) S. aemulataria (Wlk) ¢. Bog E of Big Indian L., Halifax watershed area, Nova 
Scotia, 27 June 1963, underside; (6) S. aemulataria 9, District of Columbia (no 
date), underside. Photos by Smithsonian Institution Photographic Laboratory. 

(Natural History) from New York and “East Florida.” I hereby desig- 
nate as the lectotype the male, presumed to have been taken at Trenton 
Falls, Oneida Co., New York, and it is being so labelled. I have not 
seen this specimen, but in his description of it Walker said, “hind tibiae 
hardly incrassated.” This precludes any possibility that it refers to the 
new species herein described. Also, the type locality as restricted is al- 
most certainly too far north for this new species. 

The only recognized synonym of S. aemulataria is Macaria sectomacu- 
lata Morrison (1874: 198), based on an unstated number of specimens 
from Massachusetts and New York. I have not seen the types, but again 
it would seem certain that their source is north of the range of S. 

Semiothisa promiscuata, Ferguson, new species 
eee Tere I, 4 

Description. General coloring, pattern of upperside, and wing shape almost 
exactly as in S. aemulataria, although whitish areas of wings appear a little more 
lustrous and translucent, and size somewhat larger, more nearly comparable to the 
northern S. ulsterata. Outer margin of forewing distinctly emarginate just behind 
apex, this concavity with a blackish, crescent-shaped lining. Upperside of forewing 
with antemedial and medial lines light brown, weak, nearly perpendicular to inner 
margin except angled basad just before costa; postmedial line parallel to these but 
expanded intermittently to form a series of dark brown to blackish spots, especially 
near middle of wing; postmedial bounded outwardly by a thin, pale line, and beyond 
this by an incomplete postmedial band of larger dark spots concentrated in two 
patches, as follows: a group of 3 large blackish spots in the middle of the wing, 
trisected by pale veins (M; and Cu.), closely adjacent to the mesial spots of the 

VoLUME 28, NUMBER 4 299 


LAN \ Yo 




Figs. 7-14. Genitalia and hind legs: (7) ¢ genitalia of S. promiscuata, Plummers 
Island, Maryland, 17 August 1971; (8) aedoeagus of same specimen; (9) ¢ genitalia 
of S. aemulataria, Raleigh, North Carolina, 29 April 1970; (10) aedoeagus of same 
specimen; (11) @ genitalia of S. promiscuata, Raleigh, North Carolina, 13 July 1969; 
(12) @ genitalia of S. aemulataria, Montgomery Co., Maryland, 25 May 1900; (13) 
right hind leg of S. aemulataria, Pluammers Island, Maryland; (14) right hind leg of 
S. promiscuata, Jackson, Mississippi. Drawings by the author. 


postmedial line, and a still more closely unified, subquadrate group of 2 or 3 spots 
at the costa, very thinly bisected by R; or trisected by Rs and M:, and browner than 
the nearly black mesial group. Upperside of hindwing with antemedial and post- 
medial lines light brown, weak, irregular, the latter marked by several blackish points 
on the veins; small dark discal spot present; outer third of hindwing evenly brownish, 
contrasting with paler medial and basal areas, and with less tendency to be banded 
with lighter and darker shades than in aemulataria. Underside whitish, dusted with 
reddish-brown scales; small discal spots on both wings; lines corresponding to those 
of upperside present or absent, often vague, irregular; however, immediately distad 
of the thin, wavy postmedial there is a much wider, straight or slightly curved and 
uninterrupted reddish-brown band on both wings, thinner but much more intensely 
colored than that of aemulataria. 

Length of forewing: Holotype male, 13 mm; other males, 12-13 mm; allotype 
female. 13 mm: other females, 12.5-15 mm. 

Head and body similar in the two species, including series of black dorsal markings 
on the abdomen and structure of antennae, palpi and legs, except that the male hind 
tibia of promiscuata (Fig. 14) is elongated and greatly swollen, forming the sheath 
for a large expansible hair tuft recessed into an almost full-length, longitudinal groove 
on its posterior side. In Fig. 14 the tuft is shown partly extruded. Male hind tibia 
of aemulataria (Fig. 13) hardly swollen at all and apparently lacking the hair tuft. 
Hind tarsus of promiscuata shorter than that of aemulataria. 

Male genitalia (Figs. 7, 8) most similar to those of aemulataria and its closest 
relatives, but differ in the following characters: ventral lobe of valve very broad and 
rounded at apex; elevated, bladelike ridge near apex on ventral surface of this lobe con- 
sisting mainly of a single component in promiscuata, of two separate components in 
aemulataria (Fig. 9); ventral margin of juxta straight or only slightly concave in 
promiscuata, clearly emarginate in aemulataria; tooth on gnathos and two spines on 
uncus slightly longer; sclerotized band on vesica (seen as a folded structure inside 
aedoeagus) about twice as large in promiscuata (Figs. 8, 10). Female genitalia 
larger than those of aemulataria; structures associated with ostial opening enlarged 
and more heavily sclerotized (Figs. 11, 12). 

Types. Holotype ¢ (Figs. 1, 2), Devil's Den State Park, Washington Co., 
Arkansas, 1 July 1966, R. W. Hodges, USNM Type No. 73059. Allotype 9°, Raleigh, 
North Carolina, 17 June 1970, H. H. Neunzig. Paratypes: 14, 19, Plummers 
Island [Montgomery Co.], Maryland, 17, 5 August 1971, D. R. Davis; 12, Glen Echo 
[Bethesda], Maryland, June 1914; 19, Lathrop, Maryland, 23 June 1955, E. C. 
Becker; 19, District of Columbia, 4 June 1902; 29 9, Raleigh, North Carolina, 1 
June 1970, 13 July 1969, H. H. Neunzig; 6¢ 6, 22 2, Morehead, Kentucky, 3, 8, 
19 July, 21, 29 August, 8 September 1962-63, T. N. Freeman; 192, Renfro Valley, 
Kentucky, 18 July 1955; 19, Valley Station, Kentucky, 29 August 1973, A. J. 
Brownell; 19, Homer Bird Sanctuary, Oldham Co., Kentucky, 6 September 1972, 
C. V. Covell, Jr.; 19, Elkhart, Illinois, “Aug. 1-7”; 19, McClellanville, South Caro- 
lina, 30 August 1973, R. B. Dominick; 19, Emory University, Georgia, “8-10-46,” 
H. V. Weems, Jr.; 19, Screven Co., Georgia, 8 July 1946, Otto Buchholz; 29 9, 
Gainesville, Florida, 14 May 1970, 9 July 1972, F. W. Mead; 16, Alachua Co., 
Florida, 8 April 1959, J. Perry; 246 6, 19, Torreya State Park, Liberty Co., Florida, 
23 May 1966, G. W. Rawson; 12, Greenville, Mississippi, “8-09,” G. Dorner; 16, 
Jackson, Hinds Co., Mississippi, 4 June 1960, Bryant Mather; 1¢, Town Bluff, Tyler 
Co., Texas, 27 March 1963, A. & M. E. Blanchard; 1¢, Conroe, Montgomery Co., 
Texas, 14 May 1967, A. & M. E. Blanchard. The type material is in the collections 
of the U.S. National Museum; the American Museum of Natural History; the Bio- 
systematics Research Institute, Canada Department of Agriculture, Ottawa; the Di- 
vision of Plant Industry of the Florida Deparment of Agriculture, Gainesville; Mr. 

VoLUME 28, NUMBER 4 301 

André Blanchard; Dr. C. V. Covell, Jr.; Dr. R. B. Dominick; Mr. C. P. Kimball; and 
Mr. Bryant Mather. 

Remarks. I have also seen 3 specimens regarded as too poor to include in the 
type series. These are as follows: 12, Montgomery Co., Virginia, 1 June 1901; 19, 
Renfro Valley, Kentucky, 25 May 1955; 19, Quincy, Gadsden Co., Florida, 8 No- 
vember 1966. 

Semiothisa promiscuata superficially resembles S. regulata (F.) of 
Central and South America, but the genitalia of the latter species are 
very different, more so than those of aemulataria or any of the closely 
related North American species. The greatly enlarged, swollen, male 
hind tibia is generally characteristic of the genus Semiothisa, and the 
members of the aemulataria group (Philobia) are unusual in not having 
the hind leg modified in this way. 


Morrison, H. K. 1874. New North American Lepidoptera. Proc. Boston Soc. 
Nat. Hist. 16: 194-203. 

Waker, F. 1861. List of the Specimens of Lepidopterous Insects in the Collection 
of the British Museum 23: 753-1020. 


Information to verify the acceptability of foodplants other than Quercus for 
Hemileuca maia Drury was given by Smith (1974, J. Lepid. Soc. 28: 142-145). 
The author mentions the successful rearing of maia on a species of Salix (willow) 
in 1972, from Albany Co., New York livestock collected on scrub oak, and supplied 
by me. That same year, using some of the ova from the egg mass sent to Capt. 
Smith, I reared maia on Salix (weeping willow). The larvae were fed on this food- 
plant from the beginning, not transferred to it after having been started on Quercus, 
as in the case of Capt. Smith’s program. My adults, too, emerged in September the 
same year, and were exceptionally large specimens. 

Irwin Leeuw, 1219 Crystal Lake Road, Cary, Illinois 60013. 




Department of Entomology and Nematology, University of Florida, 
Gainesville, Florida 32611 


Jerry A. POWELL 

Department of Entomological Sciences, University of California, 
Berkeley, California 94720 

Ethmia bipunctella (Fabricius) has been introduced from Europe into 
North America and during the past decade has become widely estab- 
lished in the northeastern United States and southeastern Canada (Powell, 
1973: 103). The earliest records are July 1964, on the St. Lawrence River 
in northern New York, and August 1964, in New Jersey at the mouth of 
the Hudson River. Subsequent published records are available from the 
Ottawa and Montreal areas in 1965, Connecticut in 1967, and central 
New York in 1970 (Powell, 1973). More recently accumulated collections 
include localities in Maryland, Pennsylvania and West Virginia, indi- 
cating that the adventive distribution is steadily expanding. 

In 1971 (after the above literature report was in press) specimens of 
E. bipunctella were sent to the U.S. National Museum of Natural History 
for identification from Maryland and Pennsylvania localities. K. C. Kim 
informed us (in litt.) that the earliest record for the latter state in the 
Pennsylvania State University collection is Chambersburg, Franklin Co., 
in July 1970 (R. R. Kline, coll.), while the U.S.N.M. has specimens from 
the same locality collected in May 1971 and from Friendship, Anne 
Arundel Co., Maryland in August 1970. Kim also reported that this alien 
moth was taken in several counties of Pennsylvania for the first time 
during 1973 (Penn. Cooperative Insect Report for 21 September 1973). 
The data are as follows: Franklin Co.—Letterkenny Army Depot, IV-19 
(3 moths), IV-27 (18), V-4 (33), V-16 (5), V-11 (6), V-20 (5), VI-11 
(7), VI-18 (3), VI-29 (1), VII-9 (9), VII-16 (3), VILE-27 (6G) ayineSse 
(11); Dauphin Co.—Harrisburg-York Internatl. Airport, VII-25 (5); Erie 
Co.—Erie Marine Terminal, VIII-9 (2); Centre Co.—State College, IX- 
16 (1), and Ferguson Township, IV-22 (1) (D. L. Beirlein, U.S.N.M.). 

1 Florida Agricultural Experiment Station Journal Series No. 5274. 

VoLUME 28, NUMBER 4 303 


re Lig 


2 ‘ 


Fig. 1. Spatial distribution of Ethmia bipunctella in eastern North America. 
Earliest known year of occurrence at each locality is given. 

Single males were taken by Heppner at two sites in northeastern West 
Virginia on 1 and 2 September 1973. The first was captured at Hawk 
Campground near Capon Springs, Hampshire County and the second 
about 21 km. north of Franklin, Pendleton County. Both appeared to 
be freshly emerged specimens. 


Fig. 2. Ethmia bipunctella, male, dorsal view (collected at Ottawa, Ontario, 
Canada, 2 August 1965 by H. F. Howden). 

Fig. 1 summarizes North American records by earliest year of capture, 
indicating the general outward movement from Atlantic Coast and St. 
Lawrence River points of possible introduction. The 1973 records from 
Pennsylvania and West Virginia show a continuing range extension west- 
ward and south along the Appalachian Mountains. Inasmuch as well- 
documented distribution changes are potentially useful in analyzing 
evolutionary phenomena, we encourage lepidopterists to be on the watch 
for colonies of this moth. The adult (Fig. 2) cannot be confused with 
any native species of the eastern Nearctic. The forewings are contrast- 
ingly black and white and the abdomen is bright ochreous. When at 
rest the moths are about 12-15 mm in length; spread specimens range 
21-28 mm in expanse. Nearly all collections have been made at black- 

Ethmia bipunctella is bivoltine in Europe, flying from June to July 
and in September. In southern parts of its distribution in the Old World 
and in American colonist populations, the generation pattern is not clear. 
Capture records from Pennsylvania in particular suggest a well-defined 
spring flight from late April to late May and a sporadic emergence 
through the summer months that might involve a partial third generation. 

Foodplant records in Europe include Echium vulgare (Boraginaceae), 
an introduced weed that is widespread in North America, as well as 
members of three other Holarctic genera of Boraginaceae. Probably any 

VOLUME 28, NUMBER 4 305 

native plants of this family that grow in appropriate habitats could be 
used by adventive populations of the moth. 


We thank R. W. Hodges, ARS, U.S. National Museum of Natural His- 
tory, and K. C. Kim, Department of Entomology, Pennsylvania State 
University, for providing records from the collections of their respective 


Powe LL, J. A. 1973. A systematic monograph of New World ethmiid moths (Lepi- 
doptera: Gelechioidea). Smithson. Contr. Zool. 120. 302 p. 


This brief note reports the findings of two hairstreaks which represent notable 
additions to the Nearctic region. The first of the two is previously unlisted (Dos 
Passos 1964, Lepid. Soc. Mem. No. 1; Dos Passos 1970, J. Lepid. Soc., 24(1):26-38 ) 
and was called to my attention last fall by a colleague who received one specimen 
from Mr. Wayne Klopp from the Miami, Florida area. Photographs were subsequently 
taken and submitted to Dr. F. Martin Brown, Colorado Springs, Colorado for deter- 
mination. The butterfly was identified as Electrostrymon angelica angelica Hewit- 
son, the nominate subspecies found in Cuba. In addition to the 4 original specimens 
of this species taken by Mr. Klopp in August 1973, he located a large population 
just south of Miami in January 1974. Mr. Richard Anderson, formerly of Key West, 
Florida also found the species in some numbers in that area in the latter part of 1973. 

The second species of interest here is Chlorostrymon simaethus (Drury) collected 
by Mr. Klopp and his wife Carol on Key Largo, Florida during February 1974. Pre- 
vious distributional data for this species include only continental land areas from 
South America northward to southern portions of Texas, Arizona, and California 
(Clench 1961, in Ehrlich & Ehrlich, How to Know the Butterflies, Brown, Dubuque, 
Iowa, p. 189). It is therefore unreported from any Antillean area, the origin of 
many species taken sporadically in southern Florida. Whether this is an oversight 
in the distribution or whether it has been overlooked in that region and actually 
represents an undescribed subspecies is under investigation. The author has no neo- 
tropical C. simaethus for comparison. The Florida insect is significantly distinguish- 
able from C. simaethus sarita (Skinner) from the U.S. It is expected that other 
new and interesting species will turn up in the southern Florida area from year to 
year and that collectors should keep an eye out for them, particularly the smaller, 
less conspicuous species. (Thanks to Dr. F. M. Brown for the determination for E. 
angelica and Mr. Wayne W. Klopp for examples of both species discussed. ) 

Micuaet S. Fisner, P.O. Box 7301, Denver, Colorado 80207. 

Ep. Nore: This note, and the article by R. A. Anderson in this issue, both include 
a report of the occurrence of Electrostrymon angelica angelica in Florida. For the 
record, the manuscript of Anderson was received on 8 March 1974, and that of Fisher 
on 30 April 1974. 



101 Avenida Norte #322, San Salvador, E] Salvador 

This is the fourth article of a series dealing with what my sons and 
I have found in relation to the life cycle and natural history of Rhopalo- 
cera inhabiting the vicinity of San Salvador, capital of the republic of 
E] Salvador. The first part of the series presents the subfamily Charaxinae 
of the family Nymphalidae. It started with Prepona omphale octavia 
Frihstorfer, followed by Anaea (Zaretis) itys Cramer and Anaea (Consul) 
fabius Cramer. After the present article, another on the life cycle of 
Anaea (Memphis) morvus boisduvali Comstock will continue the series. 
We undertook these investigations with the intent of presenting the life 
cycles, the foodplants, and observations on the behavior of the early 
stages and adults of the local species of Rhopalocera. There is little of 
this information in the available literature, and this applies in particular 
to the Charaxinae of Tropical America. Comstock (1961) states, “ 
there is surprisingly little to be found in the literature concerning the 
ova, larvae and pupae of the butterflies that have been discussed.” (the 
genus Anaea). Consequently the classification of this group has been 
based exclusively on morphological characteristics of the adults, which 
is not the ideal situation as implied in the following statement by Ford 
(1945), “Any classification must take into account as many as possible 
of the external and internal structures not only of the adults but of the 
early stages.” It is our hope that our articles, and the early stages which 
we have preserved in alcohol and placed in a Museum so as to be avail- 
able for students of the groups, will help in this regard. The butterflies 
mentioned in this article were identified by Dr. Lee D. Miller of the 
Allyn Museum of Entomology, where the specimens of the early stages 
have been placed. 

Anaea (Memphis) eurypyle confusa Hall was named Anaea ryphea 
by Godman and Salvin, in 1884, but was renamed by Hall in 1929. In 
order to have an idea of the habitat of this species in this country, refer 
to the first article of the series on Prepona omphale octavia (Muyshondt, 
1973). In short, A. (M.) eurypyle confusa is a denizen of coffee planta- 
tions and their neighborhood, where it is often seen feeding on decaying 
fruits or on animal and human excreta, either in the middle of the 

VoLUME 28, NUMBER 4 307 

plantations or in the roads that cross them. Its habitat is therefore limited 
to the altitudes in which coffee is planted locally, from about 700-2000 m. 
The foodplant is widely used in wind-break barriers and as live fence 
posts in coffee plantations. 

We have bred the species for a number of years now, and the results 
have been the same with small variations. Photographs have been made 
of the eggs, the different stadia, the pupae and the adults, both male and 
female. Records of development time have been kept, and specimens 
of the early stages have been preserved in alcohol and sent to the Allyn 
Museum of Entomology. The reared material was kept during develop- 
ment in transparent plastic bags under ambiental lighting and tempera- 
ture conditions. 

Life Cycle Stages 

Ege. ‘Transluscent white with greenish tinge, about 1 mm diameter, with flattened 
base and depression at micropyle. No sculpturing noticeable at 10 magnification. 
Hatch in 5 days. 

First instar larva. Head light brown, naked, roundish, with slight cleft between 
epicrania. Body light greenish brown, naked, with annulets between segments, 2.5 
mm at emergence, around 5 mm when ready to moult. Duration 5 days. 

Second instar larva. Head light brown with rudimentary horns over epicrania, 
and several whitish tubercles scattered mostly at sides of epicrania. Black ocelli. 
Body greenish brown with rings of very tiny white tubercles, three per segment. 
Whitish tubercles along subspiracular zone. Body thicker at second abdominal seg- 
ment, tapering to first thoracic segment and to last abdominal segment. Measures 
0.9-1 cm before moulting. Duration 3-5 days. 

Third instar larva. Head brown with short black horns on epicrania. Black 
vertical lines in frontal area. Scattering of white tubercles, more prominent at sides 
of head. Body greenish brown with white tubercles as in second instar. Spiracula 
dark brown surrounded by whitish ring, the first thoracic being larger than any 
other and the eighth abdominal larger than the rest. Spiracula on second and eighth 
abdominal segments are slightly higher than the others. Body thickens from first 
thoracic segment to second abdominal segment, which is surrounded by a dark band, 
and tapers then to caudal end. Dark lateral patches at fifth and seventh abdominal 
segments. Measures 1.7-1.9 cm before moulting. Duration 4—5 days. 

Fourth instar larva. Head dark brown to black with yellowish vertical lines in 
frontal area, stubby black horns on epicrania, and many prominent yellow tubercles, 
mostly at sides of epicrania and around horns. Body as in third stadium, with dark 
band along dorsal meson, more whitish tubercles along subspiracular area and across 
caudal segments, and additional lateral dark patches at third thoracic, first and sixth 
abdominal segments. Measures 3.2—-3.4 cm before moulting. Duration 5-7 days. 

Fifth instar larva. Head greenish with jet black stubby horns and very prominent 
yellow tubercles around horns and at side of epicrania; alternate greenish and yellow 
vertical lines in frontal area, those in center reaching between horns, the rest di- 
minishing gradually to sides of head. Black ocelli contrasting with yellow bordering 
line. Body green with lighter stripes dorsally from head to caudal end, and trans- 
verse rows of whitish small tubercles; spiracula contrasting over whitish patches 
forming an irregular band subspiracularly. Body now thicker than head, and dark 
patches of fourth stadium now reddish. Scarce scattering of black tubercles notice- 


Figs. 1-7. Anaea (Memphis) eurypyle confusa Hall: (1) egg, about 1 mm; (2) 
first instar larva on perch, about 3 mm; (3) second instar larva recently moulted, 
about 6 mm; (4) fourth instar larva, about 2.5 cm; (5) fifth instar larva, about 4.5 
cm; (6) close-up of head, fifth instar; (7) fifth instar larva re-entering partially 
opened funnel, note silk padding inside. 

able mostly along subspiracular zone. Measures, before entering prepupal stage, 
45-5 cm. Duration 9-11 days. 

Prepupa. Body shortens considerably and appears thicker, loses colorations of 
fifth stadium and now all light green, with the whitish small tubercles, bigger black 
spots and spiracula prominent. Stays incurvated laterally, not hanging, for one day. 

Pupa. Light green or light brown, with yellowish ridge bordering wing cases 
and across fourth abdominal segment. Cremaster black and very elaborate at base. 
Abdomen tapers abruptly from fourth segment to cremaster, and very gradually 
towards slightly bifid head. Thoracic segments keeled dorsally. Spiracula yellowish, 
very inconspicuous. Measures about 1.5 cm long, 0.9-1 cm dorsoventrally at thickest 
point, and 0.8-0.9 laterally at widest point. Duration 8-11 days. 

Adult. Both sexes same shape, with minor variations occurring even between 
individuals of same sex. Forewing more-or-less acute at apex, the outer margin 
more-or-less concave just below the apex, then more-or-less convex to tornus, and 
inner margin straight. Hindwing rounded with short tail at vein M3, anal angle 
not pronounced and with a discolored fold at inner margin. Color follows the same 
pattern in both sexes, being more vivid in the male, and very dull in the female. 
Dorsally, dominant color orange with dark brown apically; brown extending along 
costal and outer margins, leaving elongated orange patch subapically. In the male, 

VoLUME 28, NUMBER 4 309 



Figs. 8-14. Anaea (Memphis) eurypyle confusa Hall: (8-10) pupa—ventral, 
dorsal and side view; (11) male, dorsal view; (12) female, dorsal view; (13) male, 
ventral view; (14) female, ventral view. Black bars 1 cm. 


dark brown zone has bluish reflection. On hindwings, orange covers whole surface 
except for inner margin fold that is somewhat decolored, and two lighter rounded 
spots about middle of costal margin. Row of dots alongside outer margin sub- 
marginally from tail to anal angle. Ventrally both wings, in both sexes, dark grayish 


brown. Females usually larger than males; average, from tip to tip of spread fore- 
wings, 5.5 cm in female, and 5.0 cm in male. Total developmental time from 40-45 

Natural History 

During the five years we have been observing and rearing this species 
we have seen the females lay eggs on two species of Croton ( Euphor- 
biaceae): C. reflexifolius H. B. K., and, more rarely, C. niveus Jacquin. 
These species are very similar, and are known by the common name, 
Copalchi. The most apparent difference is that the fruits of C. reflexi- 
folius are muricated, but are not in C. niveus. Both species grow to small 
tree size (about 6 m) and both are used commonly to form wind-break 
barriers in coffee plantations (due to their thick foliage) and for fence 
supports. The leaves and bark of both species are very aromatic and 
bitter, and are widely used in popular medicines as infusions against 
fevers and to aromaticize alcoholic beverages. Both species keep their 
leaves year around. 

We have found in the literature (Planchon & Collin, 1895) the follow- 
ing on C. niveus: “J. Elliot Howard a signale dans cette écorce une 
matiére ameére soluble dans ether, qui au contact du chlore et de 
lamoniaque prend une teinte vert foncé. Moench n/a pu y constater la 
présence d'un alcaloide; il en a seulement retiré une huile essentielle 
constituée par un hydrocarbure, un acide organique et un principe amer 
cristallisable, la Copalchine, soluble dans lTalcool et le chloroforme.” 
Calderon y Standley (1941) state about C. reflexifolius, “La hojas y 
frutas son muy aromaticas; las hojas utilizadas en la confeccion de al- 
gunos aguardientes; la corteza como febrifugo y remedio tonico.” 

The recently emerged larvae completely devour the egg shell and stay 
under the leaf without further eating for about one day, moving after- 
wards to the border of the leaf, usually to the tip, where they choose a 
terminal vein which they eat around and bare. Using excreta stuck with 
silk they prolong the vein and use this as a resting place while not feeding, 
the head usually pointing outward. The larvae during the first, second 
and third stadia abandon this perch only for feeding purposes. During 
the fourth stadium the larvae wander about the plant for a short time 
until they select a bigger leaf, where they form a funnel-like refuge by 
rolling the leaf with the help of silk to crawl back into. From then until 
pupation the larvae keep hiding inside this funnel, leaving it momentarily 
only for feeding, which is done at dawn and dusk. The thick and tuber- 
clad head is very effective in blocking the entrance against any predator 
or injection-parasite. The excrements are expelled through the narrow 
end of the funnel. 

VoLUME 28, NUMBER 4 311 

When ready to pupate, the larvae abandon their hiding place and 
wander about the plant until a suitable place is located. This is usually 
the underside of a leaf or twig, where they weave a silken pad to which 
they affix their annal prolegs, and stay incurvated sideways, not hanging, 
for one day during which time they expel a greenish liquid mixed with 
excreta, and then pupate. 

All through the larval stages A. (M.) eurypyle confusa seems very 
apathetic. When proded with a thin brush the larvae merely extrude 
a gland located between the prothoracic legs and emit a pungent scent. 
If the proding is continued, the larvae tum around, and make biting 

The pupae are rather stiff and make only limited lateral movements 
when molested. The color of the pupae is either light green or light 
brown regardless of environmental conditions and of sex. Both morphs 
can be found simultaneously at any time of the year. The same phe- 
nomenon occurs in other species of Anaea, as well as in other Nympha- 
lidae and Brassolidae (e.g. Dynamine spp., Opsiphanes tamarindi Felder, 
and O. cassina fabricii Bdv. (Muyshondt, 1973) ). 

The adults of A. (M.) e. confusa, both male and female, are very swift 
flyers, like most Charaxinae we have observed in this country (with the 
exceptions of A. (Consul) fabius and A. (C.) electra Westwood), pro- 
ducing while in flight a rustling noise somewhat like Hesperiidae. Only 
the females when ovipositing fly slower. The female rapidly approachs 
a Copalchi plant, and then circles around it more slowly, until alighting 
under a leaf of medium development, and depositing a single egg on 
the undersurface of it, somewhere in the middle. She then resumes the 
circling around the plant and repeats the process several times before 
flying away. We have witnessed cases in which the female has oviposited 
up to six eggs without respite, at different levels on the same plant. 
Females are usually seen ovipositing late in the morning or early in the 
afternoon. Both sexes are assiduous visitors of decaying fruits and animal 
excrements, where they feed for long periods until gorged. When this 
happens, it is rather easy to net them. We have never seen this species 
feeding at flowers. The habitat of the species is restricted to coffee planta- 
tions and neighboring ravines. That means that the species is found only 
from an altitude of about 700 m up to around 2000 m, as coffee is not 
planted in El Salvador below or over these limits. 

Up to the present we have never been able to observe this species in 
courtship or while mating; in fact, we have never observed the courtship 
and mating behavior of any Charaxinae. After so much time spent in 
the field observing this and other Charaxinae without witnessing some 


sexual activity, we must assume that members of this subfamily are very 
secretive about these behaviors. 

Females dissected three days after emergence, have no eggs in their 
abdomen. It is not unusual to collect eggs that never hatch, and at times 
some eggs produce tiny wasps (Chalcidoidea). Quite often larvae of 
this species are affected by a sort of diarrhea that kills them, or by a 
disease that softens their body tissues until they burst. 


Comstock (1961) implies that nothing has been published up to now 
relating to the life cycle and behavior of the early stages of Anaea 
(Memphis) eurypyle confusa. 

As expected, the eggs of this species resemble very closely in shape 
all the eggs of the species of Charaxinae we have been able to rear, even 
to the color (with the exception of A. (Zaretis) itys whose color is 
transluscent yellow, instead of transluscent greenish-white). Further- 
more, the shape and habits of the larvae are very similar to those of 
A. (C.) fabius, A. (C.) electra and A. (Memphis) pithyusa R. Felder; 
and the pupa is quite hard to tell from that of A. (Z.) itys, A. (C.) fabius 
and A. (C.) electra, though not resembling the pupa of other species 
classified under the Memphis group of the genus Anaea that we have 
reared, such as A. (M.) pithyusa and A. (M.) morvus boisduvali. 

The wing shape of the adults of this species shows small variations in 
both sexes, even among individuals emerged during the same month. 
The behavior, flight and habitat are like those of adults of Anaea (Mem- 
phis) pithyusa, with whom they share even the foodplant. 

Like other Charaxinae, the first three stadia of Anaea (M.) eurypyle 
confusa rely for protection on their ability to imitate portions of leaf 
tissue left alongside a bared vein, while the fourth and fifth stadia hide 
within a funnel-like construction they make in a chosen leaf, and emit a 
strong odor when molested. In the funnel, the hidden larva regurguitates 
an amount of green liquid that floods the inside of the funnel and runs 
out of both ends. As the foodplant has strong aromatic and bitter prop- 
erties, it is probable that this liquid has repellent qualities for the enemies 
of the larva, and most probably the larva itself is protected by an un- 
palatable flavor derived from the foodplant. These defense mechanisms 
have proved to be very effective against “injection-parasites” at least, 
for during the eight years we have been rearing this species in our in- 
sectary, we have not found a single case of this type of parasitism. The 
protection the species has acquired against injection-parasitism does not 
work however against “ingestion-parasites,” such as the Tachinidae that 

VoLUME 28, NUMBER 4 oles 

lay their eggs on the leaf where the larvae are feeding. The amount of 
larvae killed by Tachinidae, in our experience, reaches an estimated 40%. 
The tachinid larvae usually abandon the victim during the fifth stadium 
or just after pupation. 

The adults of A. (M.) e. confusa also exhibit a combined defense 
mechanism: rapid flight with flash-and-hide effect, caused by the orange 
coloration on the dorsal surface of the wings and the cryptic grayish- 
brown coloration on the ventral side; and this cryptic coloration that 
mimics the color of a dry leaf, rendering the adults very inconspicuous 
among vegetation (or when they are sitting on surfaces such as tree 
trunks, where they even adopt a slanted position to minimize the shadow 
they project, according to the sun situation). The only time adults are 
vulnerable to predation (if they are not protected by unpalatable prop- 
erties, as we strongly suspect), is during their feeding sessions, when 
they seem to get so engorged as to lose their habitual alertness. 

Taking as a basis the developmental time of 40-45 days under labora- 
tory conditions, this species could produce about eight generations a 
year due to the fact that the foodplant remains well covered by succulent 
leaves the year around. In fact adults and larvae of the species can be 
collected at any time of the year. 

A very vulnerable stage in the life cycle of this species appears to be 
the egg stage. For some undetermined reason a considerable number 
of eggs never hatch, and some of them produce a tiny Chalcidoidea 
(which has been sent to the U.S. Dept. of Agriculture for determination). 

As said for Prepona omphale octavia (Muyshondt, 1973), this is one 
of the few species of Rhopalocera that has derived benefits from man- 
made changes in the natural ecology, i.e. by the agumentation of the 
foodplant in coffee plantations. 


We are greatly indebted to Miguel Serrano and Stephen Steinhauser 
for sharing with us their personal observations on the adults of this 
species and for giving us free access to their technical libraries; and to 
Drs. Lee D. Miller and Theodore D. Sargent for finding time in their 
crowded schedules to read the manuscripts and give constructive criti- 
cism. Special mention must go to the enthusiasm of Albert, Jr. and Pierre, 
two members of the family, who have made most of the findings on the 
early stages of this species. Specimens of the early stages have been 
sent to Allyn Museum of Entomology. 



CALDERON, SALVADOR Y P. C. STANDLEY. 1941. Lista Preliminar de Plantas de El 
Salvador. Imprenta Nacional. 

Comstock, W. P. 1961. Butterflies of the American Tropics, the Genus Anaea 
(Lepidoptera—Nymphalidae). Amer. Mus. Nat. Hist. 

Forp, E. B. 1945. Butterflies. Collins, London. 

Muysuonpt, A. 1973. Notes on the Life Cycle and Natural History of Butter- 
flies of El Salvador. I. Prepona omphale octavia (Nymphalidae). J. Lepid. 
Soc. 27: 210-219. 

1973. Notes on the life cycle and natural history of butterflies of El 

Salvador. II. Anaea (Zaretis) itys (Nymphalidae). J. Lepid. Soc. 27: 294-302. 

. 1974. Notes on the life cycle and natural history of butterflies of El 
Salvador. III. Anaea (Consul) fabius (Nymphalidae). J. Lepid. Soc. 28: 81— 

PLancuon, G. & E. Cotuiw. 1895. Les Drogues Simples d Origine Végétale. O. 
Doin, Ed. Paris. 


I wish to thank the many persons who provided assistance to me during this last 
year of my editorship. The members of the Editorial Committee of the Journal 
were most helpful as primary reviewers of submitted manuscripts. In addition, the 
following individuals reviewed one or more manuscripts upon request: L. P. Brower, 
H. A. Freeman, D. F. Owen, D. F. Schweitzer, and A. M. Stuart. I extend my 
grateful thanks for all of this help. 

My wife, Katherine, kindly provided the cover drawing (Paonias excaecatus Smith 
& Abbot), and aided in many other ways. Susan M. Moore served as an editorial 
assistant, and helped particularly by preparing the index. Finally, I wish every success 
to my successor. 


VoLUME 28, NUMBER 4 315 


Karen, Kenya 

During the course of some fifty-odd years of intermittent field work 
in Uganda and Kenya, I have gleaned a small amount of knowledge 
regarding the foodplants of the butterflies of the two territories. 

The list, in its present form, is compiled at the request of Dr. F. H. 
Rindge of the American Museum of Natural History, New York. It 
deals with some 300 species out of the 2000 known to occur in eastern 
Africa. I hope that the list, however incomplete, will be of some use 
to students of this fascinating branch of the study of Rhopalocera of 
East Africa. 

The distribution of species is closely related to the known range of 
their foodplants. It will be noted that species which are common and 
widespread have a multiplicity of foodplants belonging to several bo- 
tanical families, thus contributing to their chance of survival despite 
the rapid changes in environment now going on in these territories as 
a result of the increase in areas under cultivation and concurrent de- 
struction of indigenous forests. 

I am indebted to the following for help in determining the botanical 
material: the Director of the Kew Herbarium, through the good services 
of the late Professor Poulton of Oxford; to Dr. P. G. Greenway, at one 
time in charge of the Herbarium at the Amani Research Station, Tan- 
zania, and later Botanist in charge of the East African Herbarium, Nai- 
robi; and to Dr. B. Verdcourt, also of the East African Herbarium. 

It is regretted that, in some instances, the material submitted was in- 
adequate for specific identification, and is here listed with a query. In 
some instances, the name supplied originally is now considered to be a 
synonym, and the corrections have been made wherever possible. 

For the majority of records, the butterfly was reared from egg to 
imago on the foodplant selected by the female parent. 


Papilio dardanus Brown, eastern subspecies. Rutaceae: Teclea simplicifolia (Engl. ) 
Verdorn (= viridis Verdorn); T. nobilis Delile; T. stuhlmanni Engler; T. villosa 
N. R. F. Tayler; Toddalia asiatica Lamarck; Vespris eugeniifolia (Engl.) Ver- 
dorn; citrus, various exotic. 


Ae} as). ine) ~~ ne} tas} Ins} ine} Ine] 

Ing} ine) 

ae} las} Ine} Inc} 

Ins} Jastins}!ns] Is} Ine} ins} Ins} Is) 

. phorcas Cramer, and subspecies. Rutaceae: Teclea simplicifolia Verdorn; T. nobilis 

Delile; T. villosa Tayler. 

. mackinnoni E. Sharpe, and subspecies. Rutaceae: Teclea simplicifolia Verdorn; 

T. nobilis Delile; T. tricarpa Engler. 

. nobilis Rogenheimer, and subspecies. Canalaceae: Warburgia ugandensis Sprague. 
. lormerei Distant. Rutaceae: Clausena anisata (Wild) Oliver; Fagaropsis sp.; 

Teclea spp. 
ophidicephalus Oberthur. Rutaceae: Clausena anisata (Wild) Oliver; C. in- 
aequalis Bentham. 

. constantinus Ward, and subspecies. Rutaceae: Clausena spp.; Teclea spp. 
. hesperus Westwood. Lauraceae: Tylostemon ugandensis (Rendle) Staf. 
. rex Oberthur, and subspecies. Rutaceae: Teclea tricocarpa Engler; T. stuhlmanni 

Engler (?). 

demodacus Esperance. Rutaceae: Caledendron capensis Thunberg; Clausena 
anisata Wild (Oliver); C. inaequalis Bentham; Fagaropsis angolensis Dale; 
citrus, exotic; Toddalia asiatica Lamarck. Anacardiaceae: Pseudospondias micro- 
carpa Engler. 

. nireus Linnaeus, and subspecies. Rutaceae: Caledendron capensis Thunberg; 

Toddalia asiatica Lamarck; Clausena spp.; citrus, various exotic. 

. bromius Doubleday, and subspecies. Rutaceae: Caledendron capensis Thunberg; 

Teclea spp. 

. magda Gifford (= brontes auct.). Rutaceae: Teclea spp. 
. teita van Someren. Rutaceae: Vespris eugeniifolia Verdorn;, Teclea spp. 
. jacksoni E. Sharpe, and subspecies. Rutaceae: Clausena anisata (Wild) Oliver; 

C. inaequalis Bentham. 

. echerioides Trimen, and subspecies. Rutaceae: Clausena inaequalis Bentham; 

Toddalia asiatica Lamarck. 

. cynorta Fabricius. Rutaceae: Clausena spp. 
. ugandae Lathy. Anacardiaceae: Pseudospondias microcarpa Engler. 
. leonidas Fabricius, and subspecies. Anonaceae: Uvaria leptocladen Oliver; Uvaria 

sp.; Anona senegalensis Perseen. Apocynaceae: Landolphia ugandensis Staph.; 
L. buchannani Engler. 

. philonoe Ward. Anonaceae: Uvaria leptocladon Oliver; U. chamae Beauvais. 
. pylades Fabricius, and subspecies. Apocynaceae: Landolphia buchannani Engler; 

L. ugandensis Staph. Anonaceae: Anona senegalensis Perseen; Anona, exotic 

. policenes Cramer. Anonaceae: Uvaria bukobensis Engler; U. chamae Beauvais. 

Apocynaceae: Landolphia buchannani Engler; L. ugandensis Staph. 

. antheus Cramer, and subspecies. Apocynaceae: Landolphia ugandensis Staph. 

Anonaceae: Anonda spp. 

. sisenna Mabille. Anonaceae: Anona senegalensis Perseen. 
. porthaon Hewitson. Anonaceae: Anona spp. 

. colonna Ward. Anonaceae: Anona spp. 

. kirbyi Hewitson. Anonaceae: Anona senegalensis Perseen. 


Leptosia marginea Mabille, wigginsi Dixey, hybrida somereni Bernardi, pseudonuptilla 

Bernardi, alcesta pseudoalcesta Bernardi, medusa immaculata Aurivillius. Cap- 
paridaceae: Capparis spp. (small scandent shrubs in forest; thorns recurved; leaves 
elongate or ovate). 

Appias sylvia Fabricius, and subspecies. Euphorbiaceae: Drypetes ugandensis Hutch- 

inson; D. gerrardi Hutchinson (= battiscombei Hutchison). 

VoLUME 28, NuMBER 4 S17. 

A. lasti Smith. Euphorbiaceae: Drypetes gerrardi Hutchison; Phyllanthus sp. indet. 
A. sabina Felder, and subspecies. Euphorbiaceae: Drypetes ugandensis Hutchinson; 
D. gerrardi Hutchinson. Capparidaceae: Ritchia fragrans (Sims) G. Don. (?). 

A. epaphia Cramer, and subspecies. Capparidaceae: Boscia salicifolia Oliver; Boscia 

Pinacopteryx eriphia Godart, and subspecies. Capparidaceae: Boscia spp. 

Belenois zochalia Boisduval, and subspecies. Capparidaceae: Capparis tomentosa 
Lamarck; C. lilacinus Gilgood; C. albersi Gilgood; C. galeata Freis.; Maerua 
triphylla Richmond (= cylindricarpa Gilgood, pubescens Gilgood); M. hoenelli 
Schweinfurth. Salvadoraceae: Salvadoria persica Linnaeus. 

B. margaritacea E. Sharpe. Capparidaceae: Maerua spp. 

B. raffrayi Oberthur, and subspecies. Capparidaceae: Capparis spp. 

B. victoria Dixey, and subspecies. Capparidaceae: Capparis tomentosa Lamarck; 

Maerua spp. 

B. calypso Druce, and subspecies. Capparidaceae: Maerua spp.; Cadaba spp. 

B. subeida Felder, and subspecies. Capparidaceae: Capparis spp. 

Anaphaeois gidica Godart. Capparidaceae: Capparis spp. Salvadoraceae: Salvadoria 
persica Linnaeus. 

A. creone Cramer. Capparidaceae: Capparis spp.; Boscia spp. 

A. aurota Fabricius. Capparidaceae: Capparis spp.; Boscia spp. 

Dixeia pigea Boisduval, and subspecies. Capparidaceae: Capparis spp. 

D. doxo Godart, and subspecies. Capparidaceae: Capparis spp. 

D. spilleri Spiller. Capparidaceae: Capparis spp. 

Pieris (Belenois) solilucis Butler. Capparidaceae: Capparis tomentosa Lamarck. 

Pontia helice johnstoni Crowley. Resedaceae: Caylusia abyssinicus (Fresnius) Fischer. 
Cruciferaceae: Crucifera spp.; Epicastrum arabicum Fischer and Meyer. 

P. glauconome Klug. Cruciferaceae: Epicastrum arabicum Fischer and Meyer. 

Catopsilia florella Fabricius. Caesalpinaceae: Cassia spp. Papilionaceae: Sesbania 

Nepheronia thallasina Boisduval. Hippocrataceae: Hippocrates obtusifolia Lessner. 

N. argyia Fabricius. Rhizophoraceae: Cassipura ruwenzorensis Alsten. 

N. bouqueti Butler. Capparidaceae: Ritchea fragrans (Sims) G. Don.; R. albersi 
Gilger. Salvadoraceae: Salvadoria persica Linnaeus. 

Eronia leda Butler. Salvadoraceae: Salvadoria persica Linnaeus. Capparidaceae: 
Capparis tomentosa Linnaeus; C. caleagnous Gilger. 

Mylothris sagala Smith, and subspecies. Loranthaceae: Loranthus spp., including 
fischeri, freisiorum, dredgei, usuinensis, etc. In the absence of Loranthus, Vis- 
cum is utilized. All are parasitic. 

. chloris Fabricius, and subspecies. Loranthaceae: most species of Loranthus. 
Santalaceae: Osyris abyssinicus Hochsteter. 

. ruppelli Karsch, and subspecies. Loranthaceae: Loranthus spp. 

. poppea Cramer, and subspecies. Loranthaceae: Loranthus spp. 

. ruandana Strand. Loranthaceae: Loranthus spp. 

. bernice rubricosta Mabille. Polygonaceae: Polygonum barbatum var. fischeri 
(= setosulum A. Richard), a swamp plant. 

Colias electo Linnaeus. Caesalpinaceae: Cassia spp.; Sesbania spp. Oxalidaceae: 

Oxalis spp. 

Terias hecabe Linnaeus, and subspecies. Mimosaceae: Albizia gummifera Smith; 
Albizia spp. 

T. brigitta Cramer, and subspecies. Caesalpinaceae: Cassia spp.; Sesbania spp. 

Colotis hetaera Gerstaecker, and subspecies. Capparidaceae: Capparis spp. 

C. regina Trimen. Capparidaceae: Capparis spp.; Boscia spp. 

SSS & 


ione Godart. Capparidaceae: Capparis spp.; Boscia spp. 

. elgonensis E. Sharpe, and subspecies. Capparidaceae: Maerua spp. 

. eris Klug. Capparidaceae: Capparis spp.; Ritchea spp. 

. evarne Klug. Capparidaceae: Capparis spp.; Maerua spp. 

. incretius Butler. Capparidaceae: Capparis spp. Salvadoraceae: Salvadoria persica 

achine Cramer. Capparidaceae: Capparis spp.; Ritchea spp. 

danae Fabricius. Capparidaceae: Capparis spp. 

antigone Butler. Capparidaceae: Capparis spp. 

antevippe Butler. Capparidaceae: Capparis spp. 

evenina Wallengren. Capparidaceae: Capparis spp. 

calais amatus Fabricius. Capparidaceae: Capparis spp. 

aurigineus Baker. Capparidaceae: Capparis spp.; Boscia spp. 

vesta Reiche. Capparidaceae: Capparis spp. 

. phisadia rothschildi E. Sharpe. Salvadoraceae: Salvadoria persica Linnaeus. 

pallene rogersi Dixey. Capparidaceae: Capparis sp. indet. 

venosa Strand. Capparidaceae: Capparis spp. 

. halimede Klug. Capparidaceae: Capparis spp. 

. pleione Klug. Capparidaceae: Capparis spp. 

. celimene Lucas. Capparidaceae: Capparis spp.; Boscia spp. 



Najas (= Euphaedra) neophron Hopffer, and subspecies. Sapindaceae: Deinbollia 
borbonica Scheffler; D. kilimanjarica Taubert. 

N. uganda Aurivillius, and subspecies. Sapindaceae: Deinbollia fulvotomentella 
Baker; Allophylus subcoriacius Baker. 

N. spatiosa Mabille. Sapindaceae: Philodiscus unijugatus Radikofer (= zambesiacus 
Baker); Paullinia pinnata Linnaeus. 

N. medon Linnaeus. Sapindaceae: Philodiscus unijugatus Radikofer (= zambesiacus 
Baker); Deinbollia fulvotomentella Baker. 

N. eleus Drury, and subspecies. Sapindaceae: Philodiscus unijugatus Radikofer. 

N. coprates Druce, and subspecies. Sapindaceae: Philodiscus spp.; Allophyllus spp. 

Euryphene mardania Fabricius, and subspecies. Palmae: various palms, including 
Phoenix reclinata Jacqlin; Hyphene thebaica Mart.; borassus palm; cultivated 

Cymothoe caenis Drury, and subspecies. Flacourtiaceae: Rawsonia usambarensis 
Schaumann; R. lucida Harvey and Sender. 

C. coranus Smith. Bignonaceae: Kigelia (aethiopica) africana Bentham; K. moosa 
Sprague; Fernandoa magnifica Seemann. 

Salamis cacta Fabricius. Urticaceae: Utera hypselidendron Wedd; U. cameroonensis. 

S. temora Felder. Acanthaceae: Paulowilhelmia sclerochiton Lindau; Mimulepsis 
spatulata; Justicea spp. 

S. parhassus Drury. Acanthaceae: Asystasia schimperi T. Anderson. 

S. anacardii Linnaeus. Acanthaceae: Asystasia schimperi T. Anderson. 

Catacroptera cloanthe Cramer. Acanthaceae: Barlesia stuhlmanni (?). 

Precis octavia Cramer, and subspecies. Labiatae: Coleus barbatus Bentham; C. um- 
brosus Vatke; Plectranthus defoliatus Hochsteter; Pycnostacys sp. 

P. westermanni Westwood. Acanthaceae: Asystasia schimperi T. Anderson. 

P. oenone Linnaeus (= clelia Cramer). Acanthaceae: Asystasia spp.; Barlesia stuhl- 

P. hierta Fabricius (= cebrene Trimen). Acanthaceae:. Barlesia stuhlmanni (?). 

VOLUME 28, NUMBER 4 319 

P. chorimene Guerin. Acanthaceae: Asystasia schimperi T. Anderson; Justicia leike- 
piensis (?). 

P. sophia Fabricius. Acanthaceae: Paulowilhelmia sclerochiton Lindau. 

Pseudacraea lucretius Cramer, and subspecies. Sapotaceae: Chrysophylum viridi- 
folium Wood (= welwichii); C. albidum G. Don.; C. gorongosanum Engler. 

P. boisduvalli Doubleday, and subspecies. Sapotaceae: Chrysophylum viridifolium 
Wood; Manilkara bagshawi Moore. 

P. eurytus Linnaeus, and subspecies. Sapotaceae: Mimosops bagshawi Moore; M. 
kummel Hochstacher; Chrysophylum spp. 

Hamanumida daedulus Fabricius. Combretaceae: Combretum spp. 

Aterica galene Brown, and subspecies. Combretaceae: Quisqualia indica (= littorea 
(Engler) Exell.). 

Charaxes jasius epijasius Reichelman. Celastraceae: Gymnosporia spp.; Elaeodendron 
spp.; Maytenus spp. Gramineae: sorghum spp. 

C. jasius saturnus Baker. Celastraceae: Elaeodendron spp. Caesalpinaceae: Afzelia 
cuanzensis Welwich; Brachystygia spiciformis Bentham; B. edulis Hutchison 
and Davy. 

. jasius harrisoni E. Sharpe. Caesalpinaceae: Brachystygia spiciformis Bentham. 

. hansali Felder, and subspecies. Salvadoraceae: Salvadoria persica Linnaeus. 

castor Cramer, and subspecies. Celastraceae: Maytenus senegalensis Exall.; 

Elaeodendron spp. 

. ansorgei Rothschild, and subspecies. Melianthaceae: Bersama abyssinica Fre- 

senius; B. paullinoides Verdcoutt. 

. phoebus Butler. Melianthaceae: Bersama abyssinicus Fresenius. 

. pollux Cramer, and subspecies. Sapindaceae: Deinbollia kilimanjarica Taubert; 
D. burbonica Schaff. Melianthaceae: Bersama abyssinicus Fresenius. Euphor- 
biaceae: Fleugea microcarpa Blume. 

C. druceanus Butler, and subspecies. Myrtaceae: Syzygium caudatus Krauss; S. 

guinense Willdenow; S. sp. undet. 

C. brutus Cramer, and subspecies. Euphorbiaceae: Fluggea microcarpa Blume. 
Melianthaceae: Bersama spp. Tiliaceae: Grewia spp. Meliaceae: Melia vol- 
kensii Gurke; M. azarach (exotic); Ekebergia capensis Sparmann. 

C. lucretius Cramer, and subspecies. Anonaceae: Anona senegalensis Perseen. 

C. violetta Smith, and subspecies. Caesalpinaceae: Afzelia cuanzensis Welwich; 
Brachystygia spiciformis Bentham; B. edulis Hutchison and Davy. Sapindaceae: 
Deinbollia kilimanjarica Taubert. 

C. pythodorus Hewitson, and subspecies. Papilionaceae: Crabia brownei Dunn; C. 
laurentii Willdenow; C. brevicaudata Dunn. 

C. etesipe Godart, and subspecies. Euphorbiaceae: Ricinus communis Linnaeus; 
Croton megalocarpa Hutchison; Phyllanthus meruensis Pax; P. guinensis Pax; 
Tragia benthamii Pax; (= cordifolia). Mimosaceae: Entada abyssinicus Rich- 
mond; E. gigas Fawcett and Randle; E. scandens (Linnaeus) Bentham. Caesal- 
pinaceae: Afzelia cuanzensis Welwich. Papilionaceae: Dalbergia microcarpa 
Taub and Baker. 

. penricei Rothschild. Mimosaceae: Entada spp. 

. achaemenes Felder. Caesalpinaceae: Brachystygia spiciformis Bentham; B. oliveri 
Fawcett and Randle; B. randii Baker; B. appendiculta Bentham. 

C. guderiana Dufrane. Caesalpinaceae: Brachystygia spiciformis Bentham; B. edulis 
Hutchison and Davy. Papilionaceae: Balbergia melanoxylon Guillemin and 

C. blanda kenya Poulton. Caesalpinaceae: Brachystygia spiciformis Bentham;  B. 
edulis Hutchison and Davy. 

C. jahlusa Trimen, and subspecies. Mimosaceae: Acacia spp. 

aa a aoe 

ip) (@) 



a Bae 

baumanni Rogenhofer, and subspecies. Mimosaceae: Acacia pennata Willdenow; 
A. seval Delil; Peterolobium stellatum Brenan (= lacerans R. Bruce). 

. anticlea Drury, and subspecies. Mimosaceae: Acacia poetzi Hauman; A. pennata 


. pleione Godart, and subspecies. Mimosaceae: Acacia pennata Willdenow; Acacia 


. paphianus Westwood, and subspecies. Mimosaceae: Acacia spp. 
. manica Trimen, and subspecies. Mimosaceae: Albizia antuesiana Harmsworth. 

Papilionaceae: Dalbergia nyasae (?). 

. fulgurata Aurivillius. Caesalpinaceae: Erythrophleum africana Guillemin and 


. adubyni Poulton, and subspecies. Mimosaceae: Albizia gummifera C. A. Smith. 
. berkeleyi van Someren, and subspecies. Mimosaceae: Albizia gummifera C. A. 

Smith. Rhamnaceae: Scutia myrtina Burmann. 

= baileyi van Someren. Rhamnaceae: Scutia myrtina Burmann. 

candiope Godart. Euphorbiaceae: Croton megalocarpa Hutchison; C. macro- 
stachys Delil; C. dichogamus Pax; C. sylvaticum Krauss. 

. boueti Feisthamel, and subspecies. Gramineae: Arundinaria alpinus K. Schaumann; 

Oxytenanthera abyssinica Munro. 

. lasti Smith. Caesalpinaceae: Afzelia cuanzensis Welwich; Paramacrolobium 

coeruleum (Taub) Leonard. 

. protoclea Feisthamel, and subspecies. Caesalpinaceae: Afzelia cuanzensis Wel- 

wich. Myrtaceae: Syzygium guinensis Willdenow; S. caudatum Krauss. 

. lactetinctus Karsch, and subspecies. Myrtaceae: Syzygium cordatum Krauss. 
. eudoxus Drury, and subspecies. Araliaceae: Schefflera spp. Myrtaceae: Syzygium 


. cynthia Butler, and subspecies. Guttiferae: Garcinia sp. (unconfirmed; Sevasto- 


. tiridates Cramer, and subspecies. Sapindaceae: Phialodiscus unijugatus Baker. 

Linaceae: Hugonia platysepala Oliver (?); H. castaneifolia Engler. Ulmaceae: 
Celtis africana Burmeister; C. durandi Engler; Chaetacme arisata Planch 
(= microcarpa). Tiliaceae: Grewia mollis Jussieu; G. tricocarpa Hochsteter 
(= nyanzae Drummond). Bombacaceae: Bombax reflexcum Sprague. Flacour- 
ticaea: Flacourtia indica Merrill. Malvaceae: Hibiscus sp. undet. 

. bipunctatus Rothschild. Sapindaceae: Phialodiscus unijugatus Baker. 
. numenes Hewitson, and subspecies. Linaceae: Hugonia platysepala Oliver. 

Tiliaceae: Grewia mollis Jussieu; G. forbesi Masters. Papilionaceae: Erythrina 

abyssinica Lamarck; E. excelsa Baker. Sapindaceae: Deinbollia fulvotomentella 

. bohemani Felder. Caesalpinaceae: Afzelia cuanzensis Welwich. 
. xiphares Cramer, and subspecies. Laurinaceae: Cryptocarya spp. 
. nandina Rothschild. Euphorbiaceae: Drypetes gerrardi Hutchinson (= battis- 

combei Hutchinson). Papilionaceae: Crabia brownei Dunn (= ellioti Dunn). 
cithaeron Felder, and subspecies. Sterculiaceae: Cola laurifolia Masters. Ul- 
maceae: Chaetacme cristata Planch (= microcarpa). Papilionaceae: Crabia 
brownei Dunn. Celastraceae (Hippocrataceae): Hippocrates obtusifolia. Caesal- 
pinaceae: Afzelia cuanzensis Welwich. 

. etheocles Cramer, and subspecies. Rhamnaceae: Scutia myrtina Burmann. UI- 

maceae: Celtis durandi Engler. 

. viola Baker, and subspecies. Mimosaceae: Albizia coriaria Oliver; A. adianthi- 

folia Schoumacher (= sassa, fastigiata); Entada abyssinica Richmann; E. gigas 
Fawcet and Randle; Acacia mellifera Bentham. 

. cedreatis Hewitson. Mimosaceae: Albizia grandibracteata Taub; A. brownei 

Oliver; A. zygia Macbride; A. spp. 

VoLUME 28, NUMBER 4 O21 

C. ethalion Boisduval. Mimosaceae: Parkia filicoidea Oliver; Piptadenia buchannani 
Baker; Tamarindus indicus (exotic). Rhamnaceae: Scutia myrtina Burmann; 
S. buxifolia. 

C. virilis Rothschild. Legumonosae: Adenanthera pavonina (?). 

C. alpinus van Someren, and subspecies. Mimosaceae: Albizia gummifera Smith. 
Rhamnaceae: Scutia myrtina Burmann. 

C. zingha Stoll. Linaceae: Hugonia castaneifolia Engler; H. platysepala Oliver. 

C. eupale Druce, and subspecies. Mimosaceae: Albizia gummifera Bentham; A. zygia 
Macbride. Rhamnaceae: Scutia myrtina Burmeister. 

C. dilutus Rothschild, and subspecies. Mimosaceae: Albizia gummifera (Bentham ) 

C. subornatus Schauman, and subspecies. Mimosaceae: Albizia brownei Oliver; A. 
gummifera (Bentham) Smith. 

C. zoolina Westwood, and subspecies. Mimosaceae: Acacia pennata Wildenow; 
Acacia spp. 

C. varanes Cramer, and subspecies. Sapindaceae: Allophylus macrostachys Gilger; 
A. subcoriacius Baker; A. glaucescens. 

C. fulvescens Aurivillius, and subspecies. Sapindaceae: Allophylus macrobothrys 

C. acuminatus Thurau, and subspecies. Sapindaceae: Allophylus spp. Melanthaceae: 
Bersama abyssinicus Fresenius. 

Palla ussheri Baker, and subspecies. Convolvulaceae: Bonamia poranoides Hallier 
(= Metaporana densiflora Hallier ). 

P. violinitens Crowley, and subspecies. Convolvulaceae: Bonamia poranoides Hallier 
(= Metaporana densiflora Hallier). Verbenaceae: Clerodendron kentrocaule 

Phalantha (= Atella) phalantha Drury. Samydaceae: Trimeria spp. Caelastraceae: 
Gymnosporia spp. 

P. columbina Cramer. Flacourtiaceae: Scolopia spp. 

Lachnoptera iole Fabricius. Flacourtiaceae: Rawsonia lucida Harvey and Sonder. 

L. ayersi Trimen. Flacourtiaceae: Rawsonia usambarensis Schaumann. 

Catuna crithea Drury. Melianthaceae: Bersama abyssinicus Fresenius. 

Pseudoneptis coenibita Drury. Moraceae: Antiaris toxicaria Leschenault. 

Eurytela dryope Cramer, and subspecies. Euphorbiaceae: Ricinus communis Lin- 
naeus; Tragia benthami Pax (= cordifolia). 

E. hiabas Drury, and subspecies. Euphorbiaceae: Tragia benthami Pax (= cordifolia); 
Ricinus communis Linnaeus. 

Issoria excelsior Baker. Violaceae: Viola abyssinica. 

I. hanningtoni Elwes. Violaceae: Viola abyssinica. 

Neptis seclava Boisduval. Urticaceae: Australina accuminata Welwich; Acalypha 
paniculata; Quisqualis sp. 

N. kariakofi Overlaet. Sapindaceae: Paullinia pinnata Linnaeus. Urticaceae: Pilea 

engleri Randle; Acalypha paniculata. 

. laeta Overlaet. Mimosaceae: Albizia zygia Macbride. 

. nemetes Hewitson, and _ subspecies. Euphorbiaceae: Alchornea cordifolia 


trigonophora Butler, and subspecies. Sapindaceae: Paullinia pinnata Linnaeus. 

rogersi Eltringham. Sapindaceae: Paullinia pinnata Linnaeus. 

strigata Aurivillius. Verbenaceae: Clerodendron capitatum Schaumacher. 

. poultoni Eltringham. Verbenaceae: Clerodendron spp. Sapindaceae: Paullinia 

sp., near pinnata Linnaeus. 

melicerta Drury. Euphorbiaceae: Alchornea cordifolia Schaumacher. 

. nyssiades clarei Neave. Sapindaceae: Paullinia pinnata Linnaeus. 

. lativittata Strand. Euphorbiaceae: Cycina sp. undet. 

ae ae ae ee 


Precis stygia Aurivillius, and subspecies. Acanthaceae: Paulowilhelmia sclerochiton 

P. tera elgiva Drury. Acanthaceae: Ruellia patulata Jacquin. 

P. natalica Felder. Acanthaceae: Ruellia patulata Jacquin; Asystasia coromandeliana 

P. orithya madagascariensis Guerin. Labiatiae: Englerastrum scandens _ Alston; 
Plectranthus spp. 

P. limnoria Klug. Acanthaceae: Asystasia spp. 

P. tugela Trimen. Labiatiae: Englerastrum scandens Alston. 

Vanessa cardui Linnaeus. Urticaceae: Urtica massaica Milbred; Obetia pennatifida 
Baker; Geriadina condensata Wedd; Laportia allipes Hooker. Compositae: 
Gnaphalium declinatum Lesson (= aero o9)\ Heliochrysum spp. Boraginaceae: 
Cyanoglossum caeruleum De Candolle; C. lanceolatum Forskel. Malvaceae: 
Malva verticaliata Linnaeus. 

Antanartia abyssinica Felder. Urticaceae: Urtica massaica Milbred; Obetia pennati- 
fida Baker. 

A. hippomene Hubner, and subspecies. Urticaceae: Australina acuminata Wedde- 
now; Pouzolzia parasiticus Schweinfurth. 

A. delius Drury. Urticaceae: Australina acuminata Weddenow; Pouzolzia parasiticus 

A. schoenia Trimen. Urticaceae: Pouzolzia parasiticus Schweinfurth. 

Aterica galene Brown, and subspecies. Combretaceae: Quisqualis littorea (Engler) 
Exe. (Sevastopulo ). 

Hypolimnas missipus Linnaeus. Portulacaceae: Portulaca quadrifida Linnaeus; Talium 


anthedon Doubleday. Urticaceae: Urera hypselidendron Weddenow. 

salmacis Drury. Urticaceae: Urera hypselidendron Weddenow. 

monteironis Druce. Urticaceae: Urera hypselidendron Weddenow; Urera spp. 

antevorta Distant. Urticaceae: Urera spp. 

usambarae Westwood. Urticaceae: Urera hypselidendron Weddenow. 

. dubius De Bauvais, and subspecies. Urticaceae: Urera hypselidendron Weddenow. 

Vererone garega iansclh. Euphorbiaceae: Sapium mannicum Bentham. 

A. boisduwalli Wallengren. Euphorbiaceae: Sapium mannicum Bentham. 

A. occidentalium Moore. Euphorbiaceae: Macaranga schweinfurthi Pax. 

A. morantii Trimen. Euphorbiaceae: Macaranga kilimanjarica Pax; Excoecaria bussei 



Bematistes quadricolor Rogenhofer, and subspecies. Passifloraceae: Adenia cissam- 
peloides (Planch) Harms; Vitis spp. 

B. anganice Hewitson, and subspecies. Passifloraceae: Tryphostemma zanzibaricum 
Masters; Adenia cissampeloides (Planch) Harms. 

B. poggei Doubleday, and subspecies. Passifloraceae: Adenia spp. 

B. tellus Aurivillius, and subspecies. Passifloraceae: Adenia spp. 

Acraea johnstoni Godman, and subspecies. Urticaceae: Poulzolzia parasitica (Forsk) 
Schweinfurth; cultivated New Zealand hemp (exotic). 

A. lycoa Godart, and subspecies. Urticaceae: Poulzolzia parasitica (Forsk) Schwein- 

A. esebria Hewitson, and subspecies. Urticaceae: Poulzolzia parasitica (Forsk) 

A. rabbai mombasa Smith. Passifloraceae: Tryphostemma zanzibaricum Masters. 

A. zetes Linnaeus, and subspecies. Passifloraceae: Tryphostemma zanzibaricum 
Masters; Adenia cissampeloides Harms; Adenia lobata (?). 

VoLUME 28, NUMBER 4 S25 

bs PVPS PS sf PS a Be Se Pips os 

. asboloplintha Karsch, and subspecies. Passifloraceae: Tryphostemma zanzibaricum 

Masters; Adenia lobata. 

. natalica Boisduval, and subspecies. Passifloraceae: Adenia cissampeloides Harms; 

Adenia lobata (?). 
insignis Distant. Passifloraceae: Vitis spp. 

quirinalis Smith. Urticaceae: Urera hypselidendron (Hochst) Weddenow. 
disjuncta Smith. Urticaceae: Urera hypselidendron (Hochst ) Weddenow. 

. amacitiae Heron, and subspecies. Urticaceae: Urera hypselidendron (Hochst ) 

alcippoides Le Doux. Urticaceae: Urera hypselidendron (Hochst) Weddenow. 
neobule Doubleday. Passifloraceae: Tryphostemma zanzibaricum Masters. 

. pharsalus Ward, and subspecies. Moraceae: Ficus exasperata Vahl; F. sycomorus 


. quirina Fabricius, and subspecies. Violaceae: Rinorea poggei Engler; R. conwal- 

lariflora Brandt. 

cerasa Hewitson. Flacourtiaceae: Rawsonia lucida Harvey and Sond; R. usam- 
barensis K. Schauman. Violaceae: Rinorea conwallarifolia Brandt. 

uvui Smith, and subspecies. ‘Tiliaceae: Triumfetta rhomboidea Jacquin; T. 
macrophylla Schaumann; Sparrmannia ricinocarpa O. Kuntz. 

. bonasia Fabricius, and subspecies. Tiliaceae: Triumfetta macrophylla Schaumann; 

T. ruwenzorensis Sprague. 

. acerata Hewitson. Convolvulaceae: Ipomea batatas Linnaeus. 
. rangatana Eltringham, and subspecies. Lythraceae: Nesaea pediculata; Rotola sp. 

undet. Rosaceae: Alchemilla gracilipes Engler. 

. eponina Cramer, and subspecies. Tiliaceae: Triumfetta macrophylla Schaumann; 

T. rhomboides Jacquin. 
rahira Boisduval, and subspecies. Compositae: Eregeron canadense Linnaeus. 

. aequatorialis Neave, and subspecies. Passifloraceae: Passiflora sp. undet. Mal- 

vaceae: Malva verticillata Linnaeus. 

. caecilia Fabricius, and subspecies. Passifloraceae: Adenia cissampeloides (Planch) 

acrita Hewitson, and subspecies. Passifloraceae: Adenia spp. 
doubledayi Guerin. Passifloraceae: Adenia spp. 
sykesi Sharpe. Passifloraceae: Adenia spp. 

. calderina Hewitson. Passifloraceae: Adenia cissampeloides (Planch) Harms. 

. excelsior Sharpe, and subspecies. Tiliaceae: Triumfetta macrophylla Schaumann. 
. anacreon Trimen. Rosaceae: Alchemilla gracilipes Engler. 

. cabira Hopfter. Tiliaceae: Triumfetta spp. 

. oncaea Hopfter. Ampelidaceae: Vitis spp. Passifloraceae: Adenia spp. Flacourtia- 

ceae: Oncoba routledgei Sprague. 

Pardopsis punctatissima Boisduval. Tiliaceae: Sp. indet. (Luganda name, “subi” ). 


Danaus chrysippus Linnaeus. Asclepidaceae: Gomphocarpus fructicosus Linnaeus 

(= phillipsae); G. physocarpa Meyer; G. semilunata Richard; G. kaesneri Brown; 
G. stenophyllus Oliver (= leucocarpa); Stathmostelma gigantiflorum Schaumann; 
S. pedunculatum Decnesni (= macrantha); Aspidoglossum interruptum Bullock 
(= Schizoglossum massaicum); Kanahia lasiflora Forsk; K. glaberrima; Cynan- 
chum altoscadens Schaumann; C. abyssinicum Dacnesni; Pergularia extensa (?); 
Periploca linarifolia (?); Secamone platystigma (= africana Oliver); Caraluma 


Melinda formosa mercedonia Karsch. Asclepidaceae: Periploca linearifolia (?); 

Secamone_ platystigma (= africana (Oliver) Bullock); S. micranda (?); S. 


punctulata Decnesni; S. zambesiaca (= parvifolia (Oliver) Bullock); Crypto- 
lepsis spp. 

Tirumala petiverana Doubleday and Hewitson. Asclepidaceae: Pergularia extensa 
(= Daemia extensa). 

Amauris ansorgei Sharpe, and subspecies. Asclepidaceae: Tylophora stolzii (?); T. 
anomala N. E. Brown; Mardenia racemosa (= latifolia Schaumann); Cynanchum 

A. albimaculata Baker. Asclepidaceae: Tylophora anomala N. E. Brown; T. stolzii 
(?); Mardenia racemosa (= latifolia Schaumann); M. angolensis N. E. Brown; 
Cynanchum spp. 

A. lobengula septentrionalis Poulton. Asclepidaceae: Tylophora stolzii (?); Mardenia 
racemosa (= latifolia Schaumann); M. angolensis N. E. Brown; Gymnema syl- 
vestre (Retz) Bullock. 

A. echeria Stoll, and subspecies. Asclepidaceae: Tylophora stolzii (?); mardenia 
racemosa (= latifolia Schaumann); Secamone africana (Oliver) Bullock; S. 
parvifolia (Oliver) Bullock. 

A. niavius Linnaeus, and subspecies. Asclepidaceae: Gymnema_ sylvestre (Retz) 

A. ochlea Boisduval, and subspecies. Asclepidaceae: Tylophora stolzii (?); Cynan- 
chum abyssinica Decnesi. 


Melanitis leda Drury. Gramineae: Setaria culcata. 

Gnophodes grogani Sharpe. Gramineae: broad-blade forest grasses. 

G. chelys Fabricius. Gramineae: broad-blade forest grasses. 

G. parmeno Doubleday. Gramineae: forest grasses. 

Bicyclus (= Mycalesis) safitza Hewitson, and subspecies. Gramineae: grasses. 

B. iccius Hewitson. Gramineae: grasses. 

Neocynyra spp. Gramineae: grasses. 

Henotesia spp. Gramineae: grasses. 

Physenura spp. Gramineae: grasses. 

Ypthima spp. Gramineae: grasses. 

Aphysoneura pigmentaria Karsch, and subspecies. Gramineae: Arundinaria alpinus 


Coeliades libeon Druce. Euphorbiaceae: Drypetes gerrardi Hutchison (= battis- 

combei Hutchison ). 

C. forestans Cramer. Papilionaceae: Indigofora spp.; Sesbania spp.; Crotolaria spp. 
Asclepidaceae: Mardenia senegalensis (?); M. schimperi (Dacnesi) Bullock 
(= Dregea schimperi). Combretaceae: Combretum panniculatum Ventenat. 

. pipistratus Fabricius. Malpigiaceae: Acridocarpus zanzibaricum Jussien; A. longi- 

folium (= alopcurus Sprague). 

keithloa Wallengren, and subspecies. Malpigiaceae: Acridocarpus zanzibaricum 

Jussien; A. glaucescens Engler. Connaraceae: Bryocarpus orientalis (Baill) Baker. 

. sejuncta Mabille. Malpigiaceae: Acridocarpus zanzibaricum Jussien. 

. kenya Evans. Malpigiaceae: Acridocarpus spp. 

. anchises Gerstecker. Asclepidaceae: Mardenia angolensis N. E. Brown. 

. chalybe Westwood. Asclepidaceae: sp. indet. 

. hanno Plotz. Malpigiaceae: Acridocarpus sp. indet. 

Artitropa erinnys Trimen, and subspecies. Agavaceae: Dracaena reflexa Baker; D. 

fragrans Gawl.; D. afromontana Milbred; D. nitens Baker. 

A. comus Cramer, and subspecies. Agavaceae: Dracaena ugandae (?); D. manni (?). 

AaAaqgq Ae 

VoLUME 28, NUMBER 4 325 

A. milleri Riley, and subspecies. Agavaceae: Dracaena usambarensis Engler. 

Gamia bucholzii Plotz. Palmae: Raphia mombutorum (?); R. farinifera Hyland; 
Borasus aethiopum Martin; Phoenix reclinata Jacquin. 

G. shelleyi Sharpe. Palmae: Raphia spp.; Borassus spp.; Phoenix spp. 

Zophotes cerymica Hewitson. Palmae: Raphia spp.; Cocoa nucifer. 

Z. dysmephila Trimen. Palmae: Raphia spp.; Borassus spp.; Phoenix spp. 

Zenonia zeno Trimen. Gramineae: cultivated maize and sorghums; grasses. 

Pelopidas thrax Hubner. Gramineae: grasses. 

P. fallax Gaede. Gramineae: grasses. 

Lepella lepeletier Latreille. Gramineae: grasses. 

Androdromus philander Hopffer. Sapindaceae: Philodiscus zambesiacus Radlk. 

A. neander Plotz. Caesalpinaceae: Brachystygia randii (?); B. spiciformis Bentham. 

Eretis djaelaelae Wallengren, and subspecies. Acanthaceae: Asystasia schimperi An- 

E. lugens Rogenhofer. Acanthaceae: Asystasia schimperi Anderson; A. coromandeliana 
(?); Justicia leikepiensis (?). 

Eagris notoana Wallengren. Rhamnaceae: Scutia myrtina Kurz. 

E. subadius Gray, and subspecies. Tiliaceae: Grewia similia Schultz; G. forbesi 

E. leucetia Hewitson. Anacardiaceae: Rhus vulgaris Meikle; R. villosa Oliver. 

Gomalia elma Trimen. Malvaceae: Abutilon guinense (Schumach) Baker; A. holstii. 

Gorgyra bibulus Riley, and subspecies. Euphorbiaceae: Drypetes gerrardi Hutchison. 

Spialia spio Linnaeus. Malvaceae: Sida schimperiana Hochsteter; S. cordifolia Lin- 
naeus; S. grewioides (?); S. cuneifolia Roxburgh; Hibiscus gossyphina (?). 

S. dromus Plotz. Sterculiaceae: Waltheria americana (?); Malhamia spp. 

S. mafa Trimen. Malvaceae: Sida cuneifolia Roxburgh; S. grewioides (?); S. rhom- 
bifolia Linnaeus; Hibiscus macrantha (?). 

S. zebra bifida Higgins. Sterculiaceae: Melhamia ovata Spreng; M. velutina Forskel. 

S. kituina Karsch. Malvaceae: Sida spp. 

S. confusa Higgins. Sterculiaceae: Melhamia spp. 

S. diomus Hopffer. Tiliaceae: Triumfetta macrophylla Schuman. 

Kedestes brunneistriga Plotz. Gramineae: Setaria spp. 

Acleros mackenii Trimen. Sapindaceae: Rhus coriacius. Malpigiaceae: Acridocarpus 
longifolius (?). 

Abantis paradisea Butler. Malvaceae: Hibiscus spp. 

A. meru Evans. Compositae: Vernonia jugalis Oliver and Hierman. 

Caprona canopus Trimen. Tiliaceae: Grewia similis K. Schaumann. 

Gegenes hottentota Latreille. Gramineae: grasses. 

G. letterstedti Wallengren. Gramineae: grasses. 


The following notes on the Lycaenidae are compiled from observations 
made jointly by the late T. H. E. Jackson and myself; unfortunately, 
they are very incomplete. 


Teriomima subpunctata Kirby, micra Smith. Lays eggs on lichens on bark of trees. 

Baliochila hildegarda Kirby, dubiosa Stempffer and Bennet, fragilis Stempffer and 
Bennet, minima Bowker Smith, stygia Talbot. Eggs laid among lichens on tree 

Cnodontes vansomereni Stempffer and Bennet. Eggs laid on tree trunks among 


Telipna sanguinea Plotz, consanguinea Rebel. Eggs laid on tree trunks among lichens. 

Pentila amenida Hewitson, tachyroides Dewitz, nyasana clarensis Neave. Seen resting 
on twigs and tall grass but egg laying not observed. 

Ornipholodotos muhata Dewitz. Noted laying among lichen on twigs; larvae feed 
on lichens. 

Mimacraea poultoni Neave, marshalli dohertyi Rothschild, and subspecies. Often 
noted resting on tree trunks, head downward. Eggs laid among lichens, on 
which larvae feed. 

Hewitsonia kirbyi intermedia Joicey and Talbot. Lays eggs among small lichens on 
tree trunks, often close to ground. 

Epitola kamengoensis Jackson, cercene Hewitson, catuna carpenteri Baker. Seen lay- 
ing on twigs and tree trunks among lichens; eggs sometimes laid on dead bare 
twigs. Ants in attendance. 

Iridana incredibilis Staudinger, perdita marina Talbot. Seen laying on lichen-covered 
branches. Larvae lie up in cracks in bark within a silken protective web. 
Deloneura (Ebepius) ochrascens littoralis Talbot. Lays on tree trunks with small 
lichens; also seen laying eggs on slender bare twigs. Larvae usually in cracks 

in bark. Ants in attendance. 

Alaena caissa kagera Talbot, johanna Sharpe, ngonga Jackson, subrubra Baker. Often 
seen resting on rock faces covered with lichens, but not actually seen laying; 
frequently noted resting on flowering heads of oat grass in vicinity of rock faces. 
Pupa found in crevices of rocks. 

Spalgis lemolea Druce. Lays eggs on branches of small trees which are infested 
with coccids and scales. Larva feed on young coccids, and have been tried as a 
natural control of “mealey-bugs.” 


Lachnopnema bibulus Fabricius. Lays eggs on branches of small trees on which 
there are colonies of membracids and jassids. Larvae feed on the secretions of 
these, and also on immature membracids. Crematogaster ants also present but 
association not clear. Pupae found in ants’ nests. 

Aslauga purpurascens Holland, lamborni Baker. Eggs laid on twigs of a leafy branch 
where there is a collection of membracids. Larvae appear to prey on young 
membracids and coccids, lie up under portions of raised bark. 

Virachola (= Deudorix) dinochares Smith. Eggs laid on fruits of Syzygium cardatum 
Krauss (Myrtaceae). Larvae found within fruits, feeding on kernels; pupate 
within shell of fruit. 

V. antalus Hopffer. Eggs laid on seed pods of Leguminosae, such as wild and cul- 
tivated peas and beans. The larvae feed on seeds within pods; have also been 
noted in seed pods of Acacia stenocarpa Hoschst. (Mimosaceae) and Dolichos 
lablab Linnaeus (Papilionaceae). Pupation occurs within empty pod or in cracks 
in bark in the case of Acacia. 

V. lorisona Hewitson, coffea Jackson. Eggs laid on fruits of Rubiaceae, including 
cultivated coffee. Larvae bore into berries and eat seed sections within. Pupa- 
tion occurs within empty shell. 

. vansomereni Stempffer. Eggs laid on, and larvae feed within, kernels of Agalana 
obliqua Scheilenb. (= heterophylla Gem) (Connoraceae ). 

. dariaves Hewitson. Eggs laid on, and larvae feed within, seed pods BE Brachy- 
stygia spp. (Caesalpinaceae ). 

. suk Stempffer. Lays eggs in or around galls of Acacia spp. (Mimosaceae) oc- 
cupied by Phidole ants. 

. dohertyi Baker. Similar in habits to swk. Larvae often pupate within galls. 

Si Nea oS 

VoLUME 28, NUMBER 4 397 

V. jacksoni Talbot. Although usually placed in the genus Virachola, this species lays 
eggs on leaves of Loranthus usuiensis (Loranthaceae), on which larvae feed. 

Aphnaeus (Paraphnaeus) hutchisoni Trimen, and subspecies. Eggs laid on leaves of 
hostplant, but larvae always found within galls or swellings at point of implanta- 
tion of parasitic Loranthus. These galls are found on Acacia stenocarpa Hochst., 
and Entada abyssinicus Stend. (Mimosaceae). Larvae appear to feed on woody 
substance of swellings and are associated with ants; but tunnels used by larvae, 
and in which they pupate, free of ants. 

A. orcas Drury, and subspecies. Eggs laid on leaves of Alchornea cordifolia Schau- 
mann and Thonning (Euphorbiaceae). Larva moves to edge of leaf where it 
feeds, curling edge over and attaching it with silken threads; rests within this 
tunnel, emerging to feed; tunnel enlarged as feeding progresses, and larva pupates 
within it. 

A. propinquus Holland. Similar in habits to previous species, feeding on same food- 
plant, Alchornea cordifolia Schaumann and Thonning (Euphorbiaceae). 

Spindasis nyassae Butler. Eggs laid on leaves of Acacia stenocarpa Hochst. and En- 
tada abyssinica Steude (Mimosaceae). Larvae lie up in cracks in bark, emerging 
to feed on foliage, mainly at night; are ant attended. 

S. banyoana Baker. Eggs laid on young shoots of Acacia drepanolobium Sjostedt 
(Mimosaceae). Larvae collected by the ant Phidole and taken into galls, where 
they appear to feed on inner lining of gall and on secretions of ants; pupate 
within gall. 

S. tavetensis Lathy. Eggs laid on young shoots of Acacia drepanolobium Sjostedt 
(Mimosaceae). Larvae subsequently located on or within galls, attended by 
Phidole ants. 

S. victoriae Butler. Eggs laid on young shoots of Acacia sp. indet. (Mimosaceae ). 
Larvae are subsequently located within massed dead leaves held together with 
silken threads; pupate within this mass. 

S. homeyeri fracta Stempffer. Similar in habits to above species. 

S. appeles nairobiensis Sharpe. Seen laying on leaves of Rhus villosa Oliver (= 
vulgaris Meikle) (Anacardiaceae), but larvae and pupae were not located. 
Chloroselas pseudozeritis tytleri Riley. Eggs laid on young shoots of Acacia steno- 
carpa Hochst. (Mimosaceae). Young larvae detected in massed leaflets and 
twigs, but later found in cracks in bark, where they subsequently pupate. Cre- 

matogaster ants in attendance. 

Axiocerses amanga Westwood, and subspecies. Eggs laid on leaves of Ximenia 
americanum Linnaeus (Oleaceae). Larvae feed on leaves toward the edge, 
bringing edge of leaf over to form a tunnel, held down by silken threads; 
emerge to feed on adjacent leaves at night, returning to tunnel by day. Pupate 
within these “hides.” Attended by ants of the genus Componotus. 

A. harpax perion Cramer. Eggs laid on leaflets of Acacia drepanolobium Sjostedt 
(Mimosaceae). Larvae and pupae found within galls, attended by Phidole ants. 

A. (harpax) tjoene Wallengren. Seen laying on leaflets of Brachystygia spiciformis 
Bentham (Caesalpinaceae), but larvae not located. 

Leptomyrina lara Linnaeus. Eggs laid on leaves and stems of Kalanchoe lugardi 
Bullock (Crassulaceae). Larvae eat into leaves, feeding on soft body between 
upper and lower cortex; emerge to pupate on main stem of plant or at base 
of leaf stalk. 

Myrina silenus Fabricius. Eggs laid on young shoots of various figs (Ficus ingens 
Miquel, F. hochstetteri Reichmann; Moraceae). Larvae move onto more mature 
leaves, and pupate on stem at base of a leaf. Highly cryptic. 

M. silenus ficedula Trimen. Eggs laid on various figs, including those listed above. 
After feeding on mature leaves, larvae pupate on stem, being attached by the 
flattened “tail end,” thus simulating a small fig fruit. 


M. dermaptera dermaptera Wallengren, d. nyasae Talbot. Lays on various species of 
Moraceae, showing a preference for Ficus thonningii Blume. Habits as with 
other Myrina. 

M. sharpei Baker. Noted laying on Ficus capensis Thunberg (Moraceae), but mature 
larvae not found. 

Hypolycaena phillipus Fabricius. Eggs laid on shoots of Ximena americana Linnaeus 
(Oleaceae). Larvae found on or in seed capsules, for which they forage. 
Clerodendron capense Thunberg (Verbenaceae) is also utilized. Once seen 
laying on Loranthus dredgei (Loranthaceae ). 

H. pachelica Butler. Noted laying on Combretum constrictum Lawson (Combreta- 
ceae), but other stages not found. 

Stugeta (Iolaus) bowkeri Trimen, and subspecies. Eggs laid on Ximena americana 
Linnaeus (Oleaceae) and Loranthus dredgei (Loranthaceae). Larvae on Loran- 
thus feed on leaves, and pupate on stem. 

S. marmorea olalae Stoneham. Eggs laid on, and larvae feed on, leaves of Ximena 
americana Linnaeus (Oleaceae), but all stages not located. 

S. carpenteri Stempffer. Noted laying on leaves and flowers of Loranthus fischeri 
Engler (Loranthaceae). Imagoes can be beaten out of Loranthus clumps, but 
larvae and pupae not located. 

S. mimetica Aurivillius. Eggs laid on Loranthus sp. indet. 

Argiolaus (Iolaus) crawshayi Butler, and subspecies. Eggs laid on younger leaves of 
parasitic Loranthus usuiensis Oliver and L. dredgei (Loranthaceae). Larvae 
feed on leaves and flowers, pupating on stem towards base of a leaf and at- 
tached by posterior end to a silken pad spun on surface. Pupa very cryptic. 

A. silas silarus Druce. Similar in habits to previous species and associated with 
same species of Loranthus. 

Philiolaus parasilanus Rebel, and subspecies. Associated with parasitic Loranthus sp. 
indet. ( Loranthaceae ). 

Iolaphilus ituriensis Joicey and Talbot. Lays on Loranthus sp. indet. (Loranthaceae ). 

Epamera (Iolaus) basana yalae Riley. Noted laying eggs on leaves and flowers of 
Loranthus woodfordoides Schweinfurth and L. fischeri (Engler) Balle (Loran- 
thaceae). Larvae feed largely on flowers, but eat leaves also. Pupate on stems 
of parasite or on nearby host tree. 

E. iasis albomaculatus Sharpe. Seen laying on Loranthus sp. indet. 

E. arborifera Butler. Lays on Loranthus freisiorum and L. woodfordoides Schwein- 
furth ( Loranthaceae ). 

E. tajorica Walker. Eggs laid on Loranthus (Odontella) fischeri (Engler) Balle 

( Loranthaceae ). 

E. mimosae haemus Talbot. Eggs laid on Loranthus fischeri (Engler) Balle and L. 
recurviflora (Loranthaceae ). 

Iolaus (Pseudiolaus) poultoni Riley. Seen laying on leaves and flowers of Loranthus 
recurviflora, on which larvae feed. 

Tolaus (Aphniolaus) pallene Wallengren. Eggs laid on Loranthus fischeri (Engler ) 
Balle and L. woodfordoides Schweinfurth (Loranthaceae), as well as Ximena 
americana Linnaeus (Oleaceae). 

Anthene hodsoni hodsoni Talbot, h. usamba Talbot. Eggs laid on young shoots of 
Acacia drepanolobium Sjostedt (Mimosaceae). Young larvae found in galls, 
probably taken there by attending Phidole ants. 

A. amarah Guerin. Eggs laid on Acacia sieberiana De Candolle (= purpurascens 
Vatke), as well as A. abyssinica Hochsteter and A. stenocarpa Hochsteter 
(Mimosaceae). Larvae pupate on the stems or leaves; pupa very cryptic. 
Larvae attended by Crematogaster ants. 

A. definita Butler. Common widespread species noted laying eggs on several food- 
plants of different families, including Acacia sayal Delil, A. stenocarpa Hoch- 

VOLUME 28, NuMBER 4 329 

steter, A. abyssinica Hochsteter, Albizia spp. (Mimosaceae); Bersama engleriana 
Gurke, B. abyssinica Fresenius (Melanthaceae); Kalanchoe lugardii Bullock 
(Crassulaceae); and Rhus incana Millerm (Anacardiaceae ). 

A. larydas Cramer. Eggs laid on, and larvae feed on, leaves of Albizia gummifera 
C. A. Smith, A. zygia Macbride, Acacia farnesiana Wild., and Dichrostachys 
glomerata Hutchison (Mimosaceae ). 

A. princeps Baker, ugandae Butler. Eggs laid on, and larvae feed on, leaves of 
Entada abyssinicus Richard (Mimosaceae). Pupae found on leaves and stems. 

A. livida Trimen. Eggs laid on succulent leaves and flowers of Kalanchoe crenata 
Hewitson and K. lateralis (Crassulaceae). Larvae feed on flowers by preference, 
but also on young seeds and leaves; pupate on leaves or stems. 

A. lunulata Baker. Eggs laid on, and larvae feed on, young shoots of Entada abys- 
sinica Richard (Mimosaceae). Also° noted laying on Combretum spp. (Com- 
bretaceae ). 

A. indefinita Baker. Eggs laid on young shoots of Erythrococca rigidifolia Pax (= 
bongensis Pax). Larvae move to more mature leaves and eventually pupate 
on a leaf or under cluster of dead flowers. 

A. crawshayi Butler. Foodplant is Entada abyssinica Richard. Eggs laid on tender 
leaves, on which larvae commence to feed, later moving to more mature leaves. 
Pupate on leaves or stems on bark. Adults also lay on Acacia abyssinica Hoch- 
steter (also Mimosaceae ). 

A. nigeriae Aurivillius. Lays on young leaf shoots of Acacia stenocarpa Hochsteter 
(Mimosaceae). Larvae attended by Crematogaster ants. 

A. pitmani Stempffer. Eggs laid on young shoots of Acacia stenocarpa Hochsteter, 
as well as on A. abyssinica Hochsteter and occasionally on young plants of A. 
lahai Stendel (Mimosaceae). Newly hatched larvae feed on young foliage, then 
move onto more mature leaves. Crematogaster ants in attendance. 

A. otacilla kikuyu Baker. Seen laying on young shoots of Acacia stenocarpa Hoch- 
steter and A. lahai Stendel (Mimosaceae). As young larva grows, moves from 
young leaves to mature ones. Pupa found on stems or in cracks in bark. At- 
tended by Crematogaster ants. 

Uranothauma falkensteini Dewitz. Eggs laid on young leaf shoots of Albizia adi- 
anthifolia Schauman (= fatgiata and sassa auct.) and Acacia abyssinica Hoch- 
steter (Mimosaceae). Larvae feed on young leaves, then move to mature ones, 
on which they may pupate, although usually on stem. 

U. delatorum Heron. Lays on young leaves of Albizia gummifera Smith, but larvae 
soon move onto more mature leaves and may pupate either on old leaves or on 
main stem of branch. 

U. nubifer Trimen. Prefers laying on Albizia gummifera Smith and A. coriaria Oliver, 
but may also lay on Acacia abyssinica Hochsteter (Mimosaceae ). 

Phylaria cyara Hewitson, and subspecies. Lays on tender leaf shoots of Albizia 
gummifera Smith (Mimosaceae). Larvae move onto mature leaves and either 
pupate on them or on main branch of leaf spray. 

P. heritsia Hewitson. Eggs laid on young leaves of Bridlea macrantha Baillon (Eu- 
phorbiaceae ), preferring the foliage of saplings. 

Castalius calice Hopffer. Eggs laid on young leaf shoots of Zizyphus jujubae Lin- 
naeus (= mauritiana Lamarck) (Rhamnaceae). 

C. margaritacea Sharpe, and subspecies. Seen laying on leaves of Gounia longispicata 
Engler. Pupates towards base of mature leaves. 

C. cretosa Butler. Foodplant is Zizyphus jujubae Linnaeus (Rhamnaceae). Larvae 
first eat young leaves, then move to mature ones, eating cuticle only. Pupate on 
underside of old leaves or on leaf stalks. 

C. hintza Trimen. Lays on younger leaves of Zizyphus jujubae Linnaeus (= mauri- 


taniana Lamarck) (Rhamnaceae). As larvae mature, move onto older leaves, 
but eat only outer cortex. Pupate on underside of old leaves. 

Turacus meditteraneus Baker. Eggs laid on younger leaves of Zizyphus jujubae Lin- 
naeus (= mauritaniana Lamarck) (Rhamnaceae). Larvae move from these to 
older leaves. Pupate among leaves or on stems. 

T. ungemachi Stempffer, grammicus Smith. Both species have habits similar to 
previous ones. Also lay eggs on Zizyphus abyssinica A. Richard. 

Azanus mirza Plotz. Noted laying on Allophylus alnifolius Radlikofer (Sapindaceae ) 
and Dichrostachys glomerata Hutchison (Mimosaceae ). 

A. natalensis Trimen. Eggs laid on young shoots of Acacia abyssinica Hochsteter 
(Mimosaceae). Larvae ant-attended. 

A. jeseus Guerin. Eggs laid on young shoots of Acacia stenocarpa Hochsteter and 
A. abyssinica Hochsteter (Mimosaceae). Larvae move to older foliage and 
pupate on stem of branch. 

A. isis Drury. Foodplant is Dichrostachys glomerata Hutchison. Eggs laid on young 
leaf shoots on which young larvae feed, later moving to mature foliage; even- 
tually pupating on stem of branch. 

Syntarucus telicanus Lang. Wide range of foodplants, mostly Papilionaceae: Croto- 
laria agatifolia Schweinfurth; Sesbania aegyptiaca Person; S. sesban Linnaeus; 
Indigophora tinctoria; I. erecta Hochsteter; also wild and cultivated peas and 
beans. Also feeds on Plumbago spp. (Plumbagataceae). Eggs usually laid on 
or near flowers and younger shoots. Larvae burrow into seed pods to eat seeds; 
often pupate within pods. 

Lampides baeticus Linnaeus. Lays on several species of Crotolaria (Papilionaceae ); 
oviposition on or near flowers. Larva burrows into seed pod, where it destroys 
seeds. Often noted on cultivated peas. 

Caclirius crawshayimus Aurivillius. Lays eggs on main stem of Cyanoglossum lanceo- 
latum and C. coeruleum, both wild and cultivated (Boraginaceae). Eggs de- 
posited just above ground level. Larva eats outer cortex of root, then works up 
stem, where it pupates. Imago can only emerge when stem breaks on withering. 

Cacyreus lingeus Cramer. Labiates seem the foodplants of preference; among the 
many are Coleus lactiflorus Vatk.; C. forskelli Wild.; Calamintha simensis 
Bentham; C. elgonensis Bullock (Labiatae). Eggs laid on flowers and young 
seed pods. 

Euchrysops malathana Boisduval. Eggs laid in flowering heads of Vigna monophyla 
Taubert, and other species (Papilionaceae). Larvae eat flowers and young de- 
veloping pods. 

E. osiris Hopffer, dolorosa Trimen. Both are similar in habits to malathana, utilizing 
same foodplants, mainly Vigna monophyla Taubert. 

Cupidopsis cissus Godart. Foodplant of preference seems to be Eriosema cordifolia 
Hochsteter (Papilionaceae). Eggs laid on flowering heads, and larvae burrow 
into pods, eating the seeds. 

Eicochrysops mahalakoena Wallengren. Noted laying eggs on flowers of Acacia 
saval Delil (Mimosaceae ). 

Chiliades kedonga Smith. Noted laying eggs on young shoots and flowers of Acacia 
spp. (Mimosaceae ). 

Zizula hylas Fabricius, gaika Trimen. Both species lay eggs on flowers of Oxalis 
corniculata Linnaeus (Oxalidaceae). Larvae enter seed heads. 

Actizera lucida Baker, stellata Trimen. These seen laying eggs on flower heads of 
Oxalis corniculata Linnaeus (Oxalidaceae) and Vigna spp. (Papilionaceae). 
Lepidochrysops spp. Noted that many species of this genus lay eggs on small 
Labiatae in vicinity of “harvester” ants and allied species. Young larvae collected 
by ants and taken to nest where tended, feeding on fungi (?). Imagoes emerge 

from ants’ nests after onset of the rains. 

VoLUME 28, NUMBER 4 Sol 

Lycaena abbotti Holland, phlaeus aethiopica Poulton, pseudophlaeus Lucas. These 
three species lay eggs on species of Rumex, in particular nipalensis (= bequartii 
Wild.) and abyssinicus (Polygonaceae). Eggs laid on flowering heads and 
seeds, on which young larvae feed, before moving to leaves. 

Alocides conradsi Aurivillius, ochrascens Joicey and Talbot. Females seen laying eggs 
in trails of ants. When larvae hatch, collected and taken by ants to nest. What 
they feed on not known. 

Capys catharus Riley. Lays eggs on flowers of Protea. Larvae penetrate to base of 
flower head, and feed and pupate in this area. 


During 1973 three light traps were operated at Edgard, St. John the Baptist Parish, 
Louisiana. During this period 607 butterflies, representing 28 species, were taken 
in the traps. The distribution of individuals by species in order of abundance was 
as follows: 

Asterocampa celtis celtis Panoquina ocola (Edwards ) 6 
(Boisduval & Le Conte) 191 Eurema lisa Boisduval & Le Conte 5 
Libytheana bachmanii bachmanii | Precis coenia (Hubner ) 5 
( Kirtland ) 167 Euptoieta claudia (Cramer ) 4 
Asterocampa clyton clyton Limenitis arthemis astyanax 
(Boisduval & Le Conte) 98 (Fabricius ) 4 
Euptychia hermes sosybia Panthiades m-album 
( Fabricius ) If (Boisduval & Le Conte ) 3 
Satyrium calanus falacer Epargyreus clarus clarus (Cramer) 3 
(Godart ) 15 Pyrgus communis communis 
Calycopis cercrops (Fabricius)* 13 ( Grote ) 3 
Polites vibex vibex (Geyer) 1 Colias eurytheme eurytheme 
Polygonia interrogationis Boisduval 2 
(Fabricius ) 2, Phyciodes tharos tharos (Drury) 2 
Phyciodes phaon (Edwards ) 10 Vanessa atalanta rubria 
Strymon melinus melinus Hubner 8 (Fruhstorfer ) 2 
Limenitis archippus watsoni Phoebis sennae eubule (Linnaeus) 1 
(dos Passos ) 7 Cynthia virginiensis (Drury) 1 
Hylephila phyleus (Drury) 7 Polites themistocles (Latreille ) il 
Lerema accius (Smith) 7 Atalopedes campestris (Boisduval) 1 

* Subsequent to the compilation of these data, Mr. Harry K. Clench called to the attention of 
Bryant Mather the possibility that some material from this region, previously determined as C. 
cecrops (Fabricius) might be properly assignable to C. isobeon (Butler & Druce). 

Data on occurrence and distribution of butterflies in Louisiana have been compiled 
by Lambremont (1954, Tulane Stud. Zool. 1: 125-164), Ross & Lambremont (1963, 
J. Lepid. Soc. 17: 148-158), Lambremont & Ross (1965, J. Lepid. Soc. 19: 47-52), 
Mather (1966, J. Lepid. Soc. 20: 102), and Strickland (in prep.). From a review of 
these data it appears that all of these species, except P. coenia, have not heretofore 
been recorded from St. John the Baptist Parish. Two were not known to occur in 
Louisiana at the time of Lambremont’s 1954 work or Ross & Lambremont’s of 1963, 
but were added to the Louisiana list in 1965 by Lambremont & Ross. These are S. 
calanus falacer (Godart) (reported as C. c. calanus (Hiibner)) and Panthiades m- 
album (B & L) from East Baton Rouge and West Feliciana Parishes only, respec- 

VERNON A. Brou, Jr., 6026 Vermillion Blud., New Orleans, Louisiana 70122. 


HAYWARD (1891-1972) 

F. Martin BROWN 
6715 South Marksheffel Road, Colorado Springs, Colorado 80909 

The following list of publications was compiled from a list in the 
library of the British Museum (N.H.). The original list was based upon 
Hayward’s own records and brought together by Miss N. Schechaj of 
the Instituto Miguel Lillo, Tucuman, Argentina. I have omitted several 
non-entomological articles. Three articles were in press at the time the 
original list was compiled for transmission to the British Museum (N.H.) 
in November 1972. In many cases, Hayward did not give complete 
references to his articles. Where possible I have supplied the missing 
data. In others, such as the first two, I have not been able to locate 
copies of the journal involved. 

A note on Mantis religiosa. Egyptian Gazette (1920). 

Cyprus, a holiday resort for entomologists. Egyptian Gazette (1921). 
Aporia crataegi L. in Cyprus. Entomol. 54: 212 (1921). 

Coenonympha pamphilus, var. Entomol. 54: 290 (1921). 

Some curious aberrations of Danais chrysippus L. Entomol. 55: 178-179 
(1922). [ab. axantha, p. 178; ab. candidata, p. 179, both Egypt] 

6. Danais chrysippus L. ab. candidata Hayw. Entomol. 55: 212 (1922). 

7. Hybridization in nature. Entomol. 56: 43 (1923). 

8. Polygonia c-album L. f. album Esp. in Somerset. Entomol. 56: 43 (1923). 


Mie ESO = 

Calotropis procera R. Br. as a foodplant. Entomol. 56: 43 (1923). 

Danais chrysippus L. from Upper Egypt. Proc. Entomol. Soc. London, 1923, 

15 Jb 

11. Insects at sea. Entomol. Mon. Mag. 61: 61 (1925). 

12. Notes on Egyptian Lepidoptera observed at Reservoir, Aswan, October 1919 
till) April 1922° Entomol. Rec. & J. Var., Suppl, 37: 1-16.38. 7 @le ane 

13. The butterflies of Khartoum. Entomol. 58: 112 (1925). 

14. Migration of butterflies. Entomol. 58: 147-149 (1925). 

15. Migration of butterflies. Entomol. 58: 169-170 (1925). 

16. Miscellaneous notes from Argentina, I. Introductory. Entomol. Rec. & J. Var. 
S72 WHI (1E25)). 

17. Mites in insects. Entomol. 59: 14 (1926). 

18. Miscellaneous notes from Argentina, II. Description of the larva and pupa of 
Dione vanillae L. Description of the larva and pupa of Euptoieta hortensia 
Blanch. Description of the case of Oiketicus platensis Berg. Pugnacious attitude 
of Papilio hellanichus Hew. Entomol. Rec. & J. Var. 38: 55-57 (1926). 

19. Miscellaneous notes from Argentina, III. Colias lesbia, variation in. Description 
of the larva of Precis lavinia Cram. Entomol. Rec. & J. Var. 38: 74-76 (1926). 

20. A simple breeding cage. Entomol. Rec. & J. Var. 38: 93-94 (1926). 

21. Miscellaneous notes from Argentina, IV. The scent of Catopsilia cypris. The 
egg of Lerodea eufala Edw. Entomol. Rec. & J. Var. 38: 109-110 (1926). 

22. Migration of butterflies in northeastern Argentina in 1926. Entomol. 59: 228— 

230 (1926). 

VOLUME 28, NUMBER 4 333 
















Miscellaneous notes from Argentina, V. The life history of Papilio hellanichus 
Hew. Entomol. Rec. & J. Var. 38: 116-120 (1926). 

Miscellaneous notes from Argentina, VI. The earlier stages of Papilo thoas race 
brasiliensis R. & Jord. Entomol. Rec. & J. Var. 38: 130-133 (1926). 
Miscellaneous notes from Argentina, VII. Description of the larva of Thyreion 
olivofusa Dogn. Additional descriptive notes on the larva of Thyreion olivofusa 
Dogn. Entomol. Rec. & J. Var. 39: 18-20 (1927). 

A list of insects of various orders taken at Reservoir, Aswan, Egypt, during 
1919-1922. Entomol. Rec. & J. Var. 39, Suppl. (May) (1927). 

Duration of the pupal period of Daphnis nerii. Entomol. 60: 135 (1927). 
Collecting experiences in the Rio Paranamini region of Argentina. Entomol. 
60: 163-165 (1927). 

Migration of insects in northeastern Argentina. Entomol. 60: 188-189 (1927). 
A short description of the Argentine Chaco. Trans. South London Entomol. & 
Nat. Hist. Soc. 1927-1928: 18-33. 

Miscellaneous notes from Argentina, VIII. The larva of Chloridea armigera 
Htbn. The egg and larva of Eudamus catullus Cram. Entomol. Rec. & J. 
Wang so 20=123° (1927 ). 

Miscellaneous notes from Argentina, IX. Description of the larva and pupa of 
Hamearis chilensis Fldr (Rhopalocera). Description of the larva of Anticarsia 
gemmatalis Hbn. Entomol. Rec. & J. Var. 39: 157-159 (1927). 

Daphnis nerii. Entomol. 61: 77 (1928). 

Miscellaneous notes from Argentina, X. Ants in flooded areas. Entomol. Rec. 
& J. Var. 40: 54-55 (1928). 

Migration of insects in north-eastern Argentina 1928. Entomol. 61: 210 (1928). 
Miscellaneous notes from Argentina, XI. A short description of the larva of 
Automeris aspersa Felder. Entomol. Rec. & J. Var. 41: 12-13 (1929). 

Sobre migracion de insectos con referencia especial a la Argentina. Rev. Soc. 
Entomol. Argentina 2(5): 213 (1929). 

The caterpillar and pupa of Opsiphanes invirae sub-sp. amplificataus Stich. 
Entomol. Rec. & J. Var. 41: 34-35 (1929). 

Description of the larva and pupa of Actinotis pellenea subspecies calymna 
Jord. F. zaratenis Ob. Entomol. Rec. & J. Var. 41: 76-77 (1929). 

Larval descriptions from Argentina. A short description of the larva of Maenas 
azollae Berg. Description of the larva and pupa of Pachelia ficus Linn. De- 
scription of the larva and pupa of Herse cingulata Fabr. Entomol. Rec. & J. 
Var. 41: 91-93 (1929). 

Larval descriptions from the Argentine. Description of the larva and pupa of 
Protoparce sexta subsp. parmys Cr. Entomol. Rec. & J. Var. 41: 138 (1929). 
Larval descriptions from the Argentine. The larva of Pholus labruscae L., a 
sphingid. Entomol. Rec. & J. Var. 41: 143-144 (1929). 

Description of the larva and pupa of Automeris liberia. Entomol. Rec. & J. Var. 
Al: 164-165 (1929). 

Description of the larva and pupa of Phobetron coras Cram., a limacodid from 
the Argentine. Entomol. Rec. & J. Var. 41: 180-182 (1929). 

Description of the larva of Sibine fusca Stoll, a limacodid from the Argentine. 
Entomol. Rec. & J. Var. 42: 12-13 (1930). 

Description of the larva of Ceratocampa brisotti Bdv. Entomol. Rec. & J. Var. 
42: 136-137 (1939). 

The night flight of diurnal butterflies. Entomol. News 41: 258-259 (1930). 
Description of the larva of Speocropia sp. nov. Entomol. Rec. & J. Var. 42: 
152-153 (1930). 

In far Argentina. Entomol. Rec. & J. Var. 43: 8-12 (1931). 



Some further notes on insect migration in the Argentina. Entomol. 64: 40-41 

The early stages of Eudamus undulatus. Entomol. Rec. & J. Var. 43: 36-37 
News from the Argentine. Entomol. Rec. & J. Var. 43: 77-79 (1931). 

. A great pierine migration in the Argentine. Attacks by birds and lizards on the 

migrants. Proc. Entomol. Soc. London 6: 32 (1931). 

Contributions to the lepidopterology of the Argentine. I. Entomol. Rec. & J. 
Var. 43, Suppl. (July): 1-8 (1931). 

Notas sobre una migracion de Pieris phileta automate Burm. Rev. Soc. En- 
tomol. Argentina 3: 225-232 (1931). 

Cuatro insectos abnormales. Rev. Soc. Entomol. Argentina 3: 245-246 (1931). 
Normas para describir biologias de Lepidopteros. Rev. Soc. Entomol. Argen- 
tina 3: 257-264 (1931). 

Lepidopteros Argentinos, familia Nymphalidae. Rev. Soc. Entomol. Argentina 
4; 1-200 (1931). 

Description of the larva and pupa of the Geometrid Selenis suero Guen. Rev. 
Entomol. Brasil 2: 94-97 (1932). 

Lepidopteros Argentinos, fam. Hesperidae (sic). Parte I. Subf. Pyrrhopyginae. 
Rev. Soc. Entomol. Argentina 5: 19-35 (1932). 

New forms of Pieris phileta Fabr., Lep. Pieridae. Rev. Entomol. Brasil 2: 434— 
437 (1932). 

. Lepidopteros Argentinos. Familia Hesperiidae II. Subfamilia Pyrginae “A.” 

Rev. Soc. Entomol. Argentina 5: 149-188 (1933). 

. Notas adicionales acerca de los Nymphalidos Argentinos. Rey. Soc. Entomol. 

Argentina 5: 213-218 (1933). 

Contributions to the Lepidopterology of the Argentine, II. Entomol. Rec. & J. 
Var. 45: Suppl. (July—August ): 9-11 (1933). 

Lepidopteros Argentinos. Familia Hesperiidae III. Subfamilia Pyrginae “B.” 
Rev. Soc. Entomol. Argentina 5: 219-275 (1933). 

Informe de la octava expedicion exploradora de las zonas invernales de la 
longosta (Schistocerca paranensis Burm.). Publ. Ministr. Agric. Nacional Ar- 
gentina, “Informes de las Comisiones Exploradoras, Mayo-Agosto 1933” p 
12-15, 17-19, 21-94 37, 181-204. No. 5431 (1934). 

Lepidopteros Argentinos. Familia Hesperiidae IV. Subfamilia Pamphilinae. 
Rev. Soc. Entomol. Argentina 6: 97-181 (1934). 

Lepidopteros Argentinos. Fam. Hesperiidae V. Resumen, claves, apéndices e 
indices. Rev. Soc. Entomol. Argentina 6: 183-234 (1934). 

Normas para descrivir biologias de boroboletas. _ Boletin de Agricultura, Zoo- 
tecnicae Veterinaria estado Minas Gerais, Brasil. Ser. Agric. 7(5): 267-277 

Un nuevo parasito de los citrus. Boletin de Departmento de Agricultura, Minis- 
terio de Hacienda, Prov. de Entre Rios, I(2) (1935). 

. The geographical aspects of Argentine Entomology. Entomol. Rec. & J. Var. 

47: 109-110 (1935). 

. Los Pyrginae Argentinos. Adiciones y anotaciones (Lep. Grypocera). Ann. 

Soc. Cienc. Argentina 119: 262-266 (1935). 

Pulverizacion de los citrus. Circular 1. Estacion experimental, Concordia. (Re- 
printed in various newspapers and “Rey. B.A.P. XVIII (214) 43-44.) (1935). 
Normas para sacar muestras de plantas atacadas por parasitos 0 enfermedades 
para ser examinadas. Circ. 3. Est. Exper., Concordia (1935). 

Ensayo experimental sobre el control de la sarna en las frutas citricas. Cire. 
4, Est. Exper., Concordia (1935). 

VOLUME 28, NUMBER 4 335 








Los Pamphilinae Argentinos. Adiciones y anotaciones. (Lep. Grypocera) Ann. 
Soc. Cienc. Argentina 119: 262-266 (1935). 

Notas sobre Lepidopteros (Rhop.) Argentinos con descripcion de nuevas 
especies y formas. Rev. Soc. Entomol. Argentina 7: 183-193 (1935). 
Revision de especies argentinos de Pyrrhopyginae (Lep. Hesp.). Rev. Soc. 
Entomol. Argentina 7: 123-129 (1935). 

Phyciodes liriope (Cramer). Sinonimia y distribucion, especialmente de formas 
Argentinas. Rev. Soc. Entomol. Argentina 7: 219-223 (1935). 

Hesperidos Argentinos. Notas y adiciones, con descripcion de una nueva espe- 
cies. Rev. Soc. Entomol. Argentina 7; 131-137 (1935). 

Revision de las especies argentinos del genero Actinote (Lep. Nymphal.). Rev. 
Soc. Entomol. Argentina 7: 93-97 (1935). 

Notes on Egyptian lepidoptera observed at Reservoir, Aswan, between Oct. 
1919 and April 1922. V. Pyralidae. Entomol. Rec. & J. Var. 48, Suppl. 
(August): 1-4 (1935). 

La cochinilla blanca de los citrus y su control. Cire. 9 Est. Exper., Concordia, 
4p. (1936). 

Six month collecting along the Alto Parana, Argentina. Proc. South London 
Entomol. & Nat. Hist. Soc. 1935-1936: 55-83 (1936). 

Contribucion al conocimiento de la langosta Schistocerca paranensis y sus 
enemigos naturales. Men. Anual 1934, Comisidn Central de Investigacién sobre 
la Langosta, p. 217-229 (1936). Also as Cire. 12 Est. Exper., Concordia. 
Random notes on Argentine collecting, II. An unproductive winter expedition. 
Entomol Ree, & J. Var. 48:.111-113) 117-119 (1936). 

Argentine Notes, I. Papilionidae. Entomol. Rec. & J. Var. 49: 77-80 (1937). 
Random notes on Argentine collecting, III. The ria rain forest of the Santafecino. 
Entomol. Rec. & J. Var. 49: 5-8 (1937). 

List of the Argentine species of Pholisora (Lep. Hesp.) with descriptions of 
new species. Rev. Chil. Hist. Nat. 40: 274-278 (1936). 

Hesperioidea Argentina I. Los genitales de algunas especies y adiciones a la 
bibliographia. Rev. Soc. Entomol. Argentina 8: 57-60 (“1936,” 1937). 
Hesperioidea Argentinos II. Insectos nuevos para la fauna y anotaciones sobre 
otras. Rev. Soc. Entomol. Argentina 8: 65-76 (“1936,” 1937). 

Nymphalidos Argentinos. Adiciones y anotaciones. Rev. Soc. Entomol. Argen- 
tina 8: 93-98 (“1936,” 1937). 

Description of the larva of Mallocephala deserticola Berg. Entomol. Rec. & J. 
Var. 49: 82-83 (1937). 

Some facts about beetles. Mag. Geograph. Argentina 2(7): 29-51 (1937). 

. Hesperioidea Argentina IV. Rev. Soc. Entomol. Argentina 9: 61-65 (1937). 

Hesperioidea Argentina V. Rev. Soc. Entomol. Argentina 9: 93-101 (1937). 
Dos insectos abnormales. Rev. Chil. Hist. Nat. 41: 68-70 (1937). 

Further records of insect migration in the Argentine Republic. Entomol. 75: 
6=7 (1938). 

. Pyralidae and microlepidoptera collected in Cyprus during 1920-1921. En- 

tomol. Rec. & J. Var. 50: 6-7, 29-30, 80-82, 90 (1938). 

Hesperioidea Argentina VI. Ann. Soc. Cienc. Argentina 125: 222-231 (1938). 
Some Hesperiidae from the Yungas of Bolivia. Rev. Entomol. Brasil 8: 106— 
111 (1938). 

. Hesperioidea Argentina VII. Especies y procedencias adicionales para la fauna 

Argentina. Ann. Soc. Cienc. Argentina 125: 374-383 (1938). 

. Note on a third Argentine specimen of a Colias mosaic. Entomol. Rec. & J. 

Var. 50: 79-80 (1938). 

. Edwards Myrick, 1854-1938. Rev. Soc. Entomol. Argentina 10: 87-89 (1938). 


























A new genus and new species of neotropical Hesperiidae. Rev. Entomol. Brasil 
9: 370-374 (1938). 

Hesperioidea Argentina VIII. Ann. Soc. Cienc. Argentina 124: 429-459 (1938). 
E] acaro productor del testudo, Phyllocoptera oleivorus Ashm. Almanaque, 
Ministerio de Agricultura Nacional: 341-345 (1939). 

Hesperioidea Argentina IX. Ann. Soc. Cienc. Argentina 127: 285-293 (1939). 
Hesperioidea Argentina III. Physis 17: 279-301 (1939). 

Las especies Argentinas del genero Butleria Kirby, con descripciones de dos 
nuevas especies por Brig. Gen. W. H. Evans. Physis 17: 303-310 (1939). 
Descripcion de una nueva especies de “Speocropia” Hampson (Lep. Het., Acro- 
nyctinae). Physis 17: 311-316 (1939). 

Los parasitos de los citrus y su control. Bol. Frut. y Hort. 39: 198-253 (1939). 
Ropaloceros de las Yungas de Bolivia coleccionadas por el Sr. P. Denier en 
193i ely sism ein siin—o64(GlO89))). 

Contribucion al conocimiento de las Riodinidae Argentinos. Physis 17: 317— 
374 (1939). 

Un plan para catalogar las varias especies de moscas danhinas a las frutas en 
la Argentina y establecar su dispersion geografica como base del estudio de su 
control. — Concertacion de una accion cooperativa. — Bol. Agric. y Ganaderos, 
Proy. Cordoba, No. 173: 8-9 (1939). 

Hesperioidea Argentina X. Ann. Soc. Cienc. Argentina 128: 289-296 (1939). 
New species of neotropical Hesperiidae. Rev. Entomol. Brasil 10: 517-525 

Three new Pellicia (Lep. Hesperiidae). Rev. Chil. Hist. Nat. 43: 147-151 
(“1939,” 1940). 

A new name for Hesperia fusca Reed, 1877. Rev. Chil. Hist. Nat. 43: 16 
( “1939,” 1940). 

Ninfalidos Argentinos. Notas adicionales. Ann. Soc. Cienc. Argentina 129: 
43-47 (1940). 

Distribucion de enemigos naturales de las moscas de las frutas para su control 
biologico. Cire. 79, Est. Exper. Tucuman (1940). [Reprinted: Inform. Soc. 
Rural, Concordia 4: 58 (1940); La Chacra 10(118) (1940); Rev. Ind. Agric., 
Tucuman 30: 136-138 (1940).] 

Las cochinillas. Pampa Argentina 14(155): 19 (1940). [Reprinted: El] Campo 
94(285): 41 (1940): La Chacra 11(126): 46-47 (1941).] 

Enumeracion sistematica le los Lepidopteros de Entre Rios, I. Hesperiidae. 
Mem. No. 13, Museo de Entre Rios, 20 p. (1940). 

E] pulgon verde de los cereales. Revista Indust. Agric., Tucuman 30: 176-177 
(1940). [Reprinted: Est. Exper., Tucuman, Cire. 97 (1940); Rev. Soc. Rural, 
Rosario 20( 226): 29-34; E] Campo 24(287): 12-13 (1940); La Chacra 10: 
64-65 (1940); Pampa Argentina, No. 160, p. 14-15 (1940); La Chacra 11: 26 
(1941); La Chacra 13: 70-71 (1943).] 

Hesperioidea Argentina XI. Rev. Soc. Entomol. Argentina 12: 279-297 (1940). 
A new species of Pyrrhopyge (Lep. Hesp.). Rev. Chil. Hist. Nat. 44: 33-41 
(1940). | 

La lucha contra los insectos—El papel de la entomologia en la agricultura. 
Rev. Indust. Agric., Tucuman 30: 190-194. [Reprinted: Pampa Argentina 14: 
10-11 (1940); El Campo 25(292) p. 50-52 (1941).] 

Hesperioidea Argentina XII. Ann. Soc. Cienc. Argentina 130: 70-94 (1940). 
Migration of Colias lesbia in the Argentine in 1940. Entomol. 73: 222-224 

Contribucion a la bibliografia sobre las moscas de las frutas. Boletin 31, Est. 
Exper., Tucuman (1940). 



















Lutte biologique contra la mouche des fruits. Rev. Francais del “Oranger 10 
(107) (1940). 

New species of neotropical Hesperiidae from Ecuador. Rey. Entomol. Brasil 
11: 861-877 (1940). 

Lucha biologica contra la moscas de la frutas. Dispositivo que permite la 
salida de los parasitos beneficiosos del pozo donde se arrojan la fruta atacada. 
Cire. 95, Est. Exper., Tucuman (1940) [Reprinted: Bol. Agric., Dir. Indust., 
Fom. y Agric., Mendoza 9: 70-74 (1941); Rev. Indust. Agric., Tucuman 30: 
930-233 (1940); Rev. B. A. P. 24(284): 23-27 (1941); Pampa Argentina 
15(164): 22-93 (1940).] 

Instrucciones para la recoleccion y envio de muestras vegetales y animales para 
su examen. Circ. 90, Est. Exper., Tucuman. [Reprinted: Rev. Indust. Agric., 
Tucuman 30: 180-182 (1940).] 

La Largata rosada del algodonero (Pectinophora gossypiella Saunders). Cire. 
93, Est. Exper., Tucuman (1940). [Reprinted: Rev. Indust. Agric., Tucuman 
30: 183-193 (1940); Pampa Argentina 15(166) (1941).] 

La polilla negra del duraznero, Cydia molesta (Busck.) Circ. 99, Est. Exper., 
Tucuman (1941). [Reprinted: Rev. Indust. Agric., Tucuman 31: 316-323 
(94) Sl Gampo: 25(300): 18-21 (1941); La Chacra 12(133): 68—69, 80 
(1941); Suelo Argentino 2(16): 359-361 (1943).] 

Informe anual del Depto. de Entomologia de la Estacion Experimental Agri- 
cola de Tucuman. Rev. Indust. Agric., Tucuman 31: 50-58 (1941). 

Plantas alimenticias de hesperidos argentinos. Rev. Entomol. Argentina 11: 
31-36 (1941). 

Las cochinillas de los citricos Tucumanos y su control. Boletin 32, Est. Exper., 
Tucuman (1941). [Reprinted: Suelo Argentino 2(15): 172-177 (1943).] 
Insectos de importancia economica en la zona de Concordia, Entre Rios. Rev. 
Soc. Entomol. Argentina 9: 68-109 (1941). [Reprinted: Circ. 18, Est. Exper., 
Concordia (1941).] 

Further new species of neotropical Hesperiidae from Ecuador. Rev. Entomol. 
Basia 2pe 52531. (GAT ):. 

La lucha contra moscas de las frutas. Breve resena sobre los cebos ensayados 
en diversas partes del mundo y su aplicacion. Rev. Indust. Agric., Tucuman 
ol: 331-349 (1941). 

Algunas observaciones sobre las moscas de las frutas en la Argentina. Rev. 
Indust. Agric., Tucuman 31: 324-330 (1941). 

Hesperiidarum Argentinarum Catalogus. Revista del Museo La Plata, (NS) 
2: 227-340 (1941). 

La Cucaracha de la casa y su control. Circ. 105, Est. Exper., Tucuman (1941). 
[Reprinted: Rev. Indust. Agric., Tucuman 31: 422 (1941); Suelo Argentino 
1: 762-763 (1942).] 

La folilla de la papa y su control. (Gnorimischema operculella Kell.). Circ. 
108, Est. Exper., Tucuman (1942). [Reprinted: Rev. Indust. Agric., Tucuman 
32: 4-6; Pampa Argentina 17(188): 10-11, 22 (1943).] 

Informe anual del Depto. de Entomologia, 1941. Rev. Indust. Agric., Tucu- 
man 32: 45-55 (1942). 

Hesperioidea Argentina, XIII. Rev. Soc. Cienc. Argentina 134: 64-71 (1942). 
El gusano chupador de la Cana de Azucar Diatraea saccharalis (Fabricius) en 
Tucuman. Boletin 38, Est. Exper., Tucuman. [Reprinted: La Industria Azu- 
carera 49: 256-263 (1943).] 

Nuevas especies de Hespéridos brasilehos. Papéis Avulsos, Dept. Zoologia, Sao 
Rawlon2:) 171=177 (1943). ~ 

Notes on the Hesperiidae in the collection of the Museu Nacional do Brasil. 
I. Pyrrhopyginae. Bol. Mus. Nac. Brasil 14-16: 62-82 (1942). 













7 dk 





El gusano chupador de la cafia de azucar. Cire. 115, Est. Exper., Tucuman 
(1943). [Reprinted: Rev. Indust. Agric., Tucuman 32: 315-325 (1942) (?!).] 
La polilla taladradora de la cafa de azucar, Elasmopalpus lignosellus (Zeller). 
Boletin 40, Est. Exper., Tucuman (1943). [Also: Rev. Indust. Agric. Tucu- 
man 32: 326-332 (1942).] 

El escarabajo o cascarudo rhinocerante, Strategus validus (Fabricius) y la 
cana de azucar. Circ. 116, Est. Exper., Tucuman (1943). [Also: Rev. Indust. 
Agric., Tucuman 32: 333 (1942).] 

Primera lista de insectos tucumanos perjudiciales. Pub. Misc. No. 1, Est. 
Exper. Tucuman. 110)p. (“1942.7 1943). 

La oruga de la hoja del algodonero, Alabama argillacea (Huber) en Tucuman. 
Boletin 41, Est. Exper., Tucuman (1943). 

El carbon de la cafia y los insectos. Circ. 123, Est. Exper., Tucuman (1943). 
[Reprinted: La Industria Azucarera 49: 17-18 (1944); Suel Argentino 3: 133 
(1944); El Campo 29(346): 30 (1945); Azucar 1(4): 36 (1944); Rev. Indust. 
Agric., Tucuman 34: 96-97 (1944).] 

Notas Entomo-biologicas y otras (por Rodolfo Schreiter) preparadas para su 
publicacion por K. J. Hayward. Acta Lilloana, Tucuman 1: 5-44 (1943). 
Estudios sobre Hespéridos neotropicales. I. Sobre la sinonomia y organos 
genitales de ciertas especies del género Dalla Mabille. Acta Lilloana 1: 45-53 

Hespéridos americanos cuyas larvas perjudician la cama de azucar. Rev. Indust. 
Agric., Tucuman 33: 11-18 (1944). 

O besuro ou cascudo rinocerante e a cana de acticar. Brasil Acticar, Rio de 
Janeiro 22: 69-74. (1943). 

A broca de cana de actcar. Brasil Acticar 22: 69-74 (1943). (NB. I have 
not been able to see this and the previous paper. Miss Schechaj’s list used 
the same citation for both. ) 

A mariposa perfuradora de cana de actcar. Brasil Acticar 22: 84-87 (1943). 
Informe Anual del Depto. Entomologia de la Estacion Experimental de Agri- 
cultura, Tucuman, 1942. Rev. Indust. Agric., Tucuman 33: 66-84 (1943). 
La cochinilla blanca de los citricos (Unaspis citri Comstock) en Tucuman. 
Cire. 124, Est. Exper., Tucuman (1944). [Reprinted: Rev. Corp. Frutas Ar- 
gentina 10(17): 7-15 (1944).] 

Contribucion a la bibliografia sobre el pulgon amarilla de la cafia de azucar 
(Sipha flava (Forbes)). Publ. Misc. 3, Est. Exper. Tucuman (1944). 

El pulgon amarillo de la cama de azucar (Silpha flava (Forbes) ). Cire. 125, 
Est. Exper. Tucuman (1944). 

Las moscas le last frutas en Tucuman. Cire. 126, Est. Exper. Tucuman (1944), 
[Reprinted: Suelo Argentino 4(39): 176-178, 183-189 (1944).] 

Two new neotropical Hesperiidae. Rev. Chil. Hist. Nat., 45: 63-68 (“1941,” 

Hesperioidea Argentina XIV. Rev. Soc. Entomol. Argentina 12: 173-180 

Lista de insectos perjudiciales. Primera suplemento. Publ. Misc. No. 4, Est. 
Exper. Tucuman (1944). 

Las moscas blancas (Aleyrodidae). Circ. 128, Est. Exper. Tucuman (1944). 
Los pulgones o afididos. Cire. 129, Est. Exper. Tucuman (1944). [Reprinted: 
Suelo Argentino 4(40): 255-256 (1944).] 

O pulgao amarelo da cana de acicar en Tucuman. Brasil Actcar 25: 316-332 

Modelo para una jaula que permite la distribucion de parasitos dentro de las 
pupas de sus huespedes. Rev. Indust. Agric., Tucuman 34: 23-26 (1945). 

VoLUME 28, NUMBER 4 339 







(with R. C. Williams) Hesperiidarum Rei Publicae Aequatoris Catalogus. Acta 
Zool. Lilloana 2: 63-246 (1944). 

Hesperiidae (Lep.) capturados em Porto Cabral durante una excursao a margem 
Paulista do Rio Paranda. Papéis Avulsos Dept. Zool., Sao Paulo 5: 197-202 

Memoria anual 1943 del Depto. de Entomologia de la Estacion Experimental 
de Tucuman. Rev. Indust. Agric., Tucuman 34: 151-165 (“1944,” 1945). 
Instructions para la caza y acondicionamiento de insects. Publ. Misc. No. 10, 
Instituto Miguel Lillo, Tucuman (1945). 

O “Carvao” da cana de actcar e os insetos. Brasil Acicar 26: 98 (1945). 
Hesperiidae (Lep.) capturados em Porto Cabral durante una segunda excursao 
a margem Paulista do Rio Parana. Papéis Avulsos, Dept. Zool., Sao Paulo 
7: 129-142 (1946). 

Memoria del Depto de Entomologia de la Estacion Experimental de Agricul- 
tura, Tucuman, 1944. Rev. Indust. Agric., Tucuman 36: 60-72 (1946). 
Argynnis paphia var. valenzina Esp. and some other butterflies of Somerset. 
Entomologist 80: 20-21 (1947). 

Hesperioidea Argentina XV. Acta Zool. Lilloana, Tucuman 3: 215-230 (“1946,” 

Una especies y forma nuevas de Hespéridos Argentinos. Acta Zool. Lilloana, 
Tucuman 3: 253-256 (“1946,” 1947). 

Las especies Argentinas de los géneros Mylon y Carrhenes. Acta Zool. Lilloana, 
Tucuman 3: 307-312 (“1946," 1947). 

Roswell Carter Williams. Rev. Soc. Entomol. Argentina 13: 344-345 (1947). 

. Hesperioidea Argentina XVI. Acta Zool. Lilloana, Tucuman 4: 4-18 (1947). 
. Algunas planta huespedes de las larvas de los Hespéridos americanos. Acta 

Zool. Lilloana, Tucuman 4: 19-54 (1947). 

Hesperioidea Argentina XVII. Acta Zool. Lilloana, Tucuman 4: 55-61 (1947). 
Una nueva especie de “Automalis’” (Lep. Het. Arctiidae). Acta Zool. Lilloana, 
Tucuman 4: 63-67 (1947). 

Nuevas especies de Hespéridos sudamericanos. Acta Zool. Lilloana, Tucuman 
4; 121-128 (1947). 

Hesperioidea Argentina XVIII. Acta Zool Lilloana, Tucuman 4: 133-144 

Catalogus Hesperiidarum Rei Publicae Colombianae. Acta Zool. Lilloana, 
Tucuman 4: 201-392 (1947). 

. Three new genera for neotropical Hesperiidae. Acta Zool. Lilloana, Tucuman 

5: 97-102 (1948). 

. Hesperioidea Argentina XIX. Acta Zool. Lilloana, Tucuman 5: 103-112 (1948). 

Nuevos especies de Hespéridos neotropicales. (Lep. Hesperiidae). Acta Zool. 
Lilloana, Tucuman 5: 175-183 (1948). 

. Insecta. Lepidoptera, fam. Hesperiidae, subfam. Pyrrhopyginae et Pyrginae. 

Tomo I. Del genera et species animalium Argentinorum. 389 p., 27 pl., some 
in color. Talleres Graficos Kraft, Buenos Aires. August 1948. 

Ninfalidos Argentinos. Modificaciones en su nomenclatura y en la lista de 
especies (Lep. Nymphalidae). Acta Zool. Lilloana, Tucuman 7: 5-26 (1949). 

. Una nueva especies y dos nuevas formas de Piéridos argentinos (Lep. Pieridae). 

Acta Zool. Lilloana, Tucuman 7: 135-147 (1949). 

Hesperioidea Argentina XX. Subfamily Hesperiinae: algunas cambios de no- 
menclatura. Acta Zool. Lilloana, Tucuman 7: 331-335 (1949). 

especies (Lep. Nymphalidae). Acta Zool. Lilloana, Tucuman 7: 5-26 (1949). 
Biological notes on some Hesperiidae of Para and the Amazon by A. Miles Moss. 
(Compilado y en su mayor parte redescrito por K. J. Hayward de anotaciones 



dejados por Miles Moss en yoder del Museo Britanico (Hist. Nat.) de Londres. ) 
Acta Zool. Lilloana, Tucuman 8: 1-80 (1949). 

Satiridos Argentinos nuevos para la ciencias (Lep. Satyridae). Acta Zool. 
Lilloana, Tucuman 8: 151-159 (1949). 

Nuevas especies y formas de Riodinidae de Argentina y Bolivia. (Lep. Rhop.). 
Acta Zool. Lilloana, Tucuman 8: 197-201 (1949). 

Nuevas especies del Lycaenidae de la Argentina. (Lep. Rhop.) Acta Zool. 
Lilloana, Tucuman 8: 567-581 (1949). 

Insecta. Lepidoptera, fam. Hesperiidae, subfam. Hesperiinae. Tomo II. Del 
genera at species animalium Argentinorum. 388 p., 26 pl., some in color. 
Talleres Grafico Kraft, Buenos Aires. December 1949. 

Un nuevo genero para Nymphalidae. Rev. Soc. Entomol. Argentina 14: 319- 
320 (1950). 

Catalogo sinonimico de los Rhopaloceros Argentinos excluyendo Hesperiidae. 
Acta Zool. Lilloana, Tucuman 9: 85-282 (preprint released in 1950, volume 
published in 1951). 

Las especies y formas Argentinas del género Adelpha (Lep. Nymphalidae) 
Acta Zool. Lilloana, Tucuman 9: 375-393 (1951) (preprint released in 1950). 
Estudios sobre Hesparidos neotropicales (Lep. Hesperiidae) II. Descripciones 
de nuevas especies. Acta Zool. Lilloana, Tucuman 9: 463-470 (1951) (pre- 
print released in 1950). 

Hesperioidea Argentina XXI. Rev. Soc. Entomol. Argentina 15: 53-56 (1950). 
Memoria anual del Instututo de Entomologia, 1949. Memoria anual de Uni- 
versidade, Tucuman (1950). 

Memoria anual del Instituto de Entomologia, 1950. Memoria anual de Uni- 
versidade, Tucuman (1951). 

Apuntas sobre bibliografia y nomenclatura para la licenciatura de zoologia. 
(Mimeographed). Instituto Miguel Lillo, Tucuman (1951). 
Paradichlorobenzene and mould. Entomol. Rec. & J. Var. 44: 183-184 (1952). 
Butterflies on wet ground. Entomol. Rec. & J. Var. 44: 216-220 (1952). 
Mas notas sobre Ninfalidos Argentinos. Acta Zool. Lilloana, Tucuman 10: 285- 
290 (1952). 

Una clava para las subfamilias, géneros y especies Argentinos de la familia 
Heliconiidae (Lep. Rhop.) Acta Zool. Lilloana, Tucuman 10: 311-313 (1952). 
Clave para los géneros y especies Argentinas de la familia Nymphalidae. Acta 
Zool. Lilloana, Tucuman 10: 401—421 (1952). 

Guia para la classificacion de las especies y formas Argentinas de la familia 
Papilionidae. Acta Zool. Lilloana, Tucuman 12: 279-330 (“1951,” 1952). 
Migration of butterflies in Argentina during the spring and summer of 1951-— 
1952. Proc. Royal Entomol. Soc. London (A) 28: 63-73 (1953). 

Memories of the years. Entomol. Rec. & J. Var. 45: 202-205 (1953). 

An entomologist in Argentina. The chaco santafesino. Entomol. Rec. & J. Var. 
Gas Due Dis) (Iles). 

An entomologist in Argentina II. Buenos Aires and La Rioja. Entomol. Rec. & 
J. Var. 65: 310-315 (1953). 

An entomologist in Argentina III. Exploring for locusts. Entomol. Rec. & J. 
Ware 659) 050-395) (1953) 

An entomologist in Argentina IV. Journey to Missiones. Entomol. Rec. & J. 
Var. 66: 12-16 (1954). 

An entomologist in Argentina V. Collecting in Missiones. Entomol. Rec. & J. 
Var. 66: 77-82 (1954). 

An entomologist in Argentina VI. Citrus entomologist. Entomol. Rec. & J. Var. 
66: 138-142 (1954). 

VoLUME 28, NUMBER 4 341 







Catalogo sinonimico de Ropaloceros Argentinos excluyendo Hespéridos. (Primero 
suplemento). Acta Zool. Lilloana, Tucuman 14: 353-374 (1953). 

Memoria anual 1953 del Departmento de Zoologia del Instituto Miguel Lillo. 
Publ. Fund, Miguel Lillo, Tucuman (1954). 

An entomologist in Argentina VII. Agricultural entomologist. Entomol. Rec. 
& J. Var. 66: 191-195 (1954). 

Apuntas para un curso de entomologia para la licenciatura de tecnologia azu- 
carera. Publ. Asoc. Gremial Univ. Farm. Azuc. Bioquim., Univ. Nacional, 
Tucuman, 77 p., 17 pl. (1954). 

An entomologist in Argentina VIII. Missiones revisited. Preparation, journey 
and arrival. Entomol. Rec. & J. Var. 66: 22-226 (1954). 

An entomologist in Argentina IX. Collecting in northwestern Missiones. En- 
tomol. Rec. & J. Var. 66: 247-252 (1954). 

An entomologist in Argentina X. The chaco santafesina revisited. Entomol. 

Rec. & J. Var. 66; 266-271 (1954). 

An entomologist in Argentina XI. Bahia Blanca and the Rio Negro valley. 
Entomol. Rec. & J. Var. 66: 290-293 (1954); 67: 19-20 (1955). 

Satiridos argentinos (Lep. Rhop. Satyridae) I. Los géneros (excluidos Euptychia 
y Neomaniola). Acta Zool. Lilloana, Tucuman 13: 5-66 (1953). 

Las fechas de publicacion de “Die Auslandische Schmetterlinge” de Esper. 
Acta Zool. Lilloana, Tucuman 13: 67-68 (1953). 

Tres nuevas especies de satiridos Argetinos. Rev. Soc. Entomol. Argentina 
17: 15-18 (1954). 

An entomologist in Argentina XII. Neuquen. Entomol. Rec. & J. Var. 67: 
GSI ( 1955)).. 

An entomologist in Argentina XIII. The Calchaqui Valleys. Entomol. Rec. & 
J. Var. 67: 165-169 (1955). 

Migration of butterflies in Argentina, 1953-1954. Proc. Royal Entomol. Soc. 
London (A) 30: 59-62 (1955). 

An entomologist in Argentina XIV. Mendoza. Entomol. Rec. & J. Var. 67: 
226— --- (1955). 

An entomologist in Argentina XV. Short trip to La Rioja. Entomol. Rec. & J. 
Ser T 2662270 (1955). 

The identity of Pamphila kirbyi Reed. Entomol. 88: 261 (1954). 

Nuevas Euptychia de Bolivia. Rev. Chil. Entomol. 5: 107-121 (1957). 
Cursillo de entomologia para azucareros. Publ. misc. No. 18, Inst. Miguel 
Lillo, 112 p. (published by Univ. Nacional, Tucuman). 

Migration of butterflies in Argentina (Summer 1956-1957). Entomol. 91: 
163-164 (1958). 

Satiridos Argentinos, II. Los generos (con't). Acta Zool. Lilloana, Tucumaén 
15: 161-181 (1958). (Miss Schechaj dates this 1959. ) 

Satiridos Argentinos, III. Guia para su classificacion. Acta Zool. Lilloana, 
Tucuman 15: 199-296 (1958). 

Catalogo sinonimico de los Ropaloceros Argentinos excluyendo Hesperidae. 
(Segundo suplemento.) Acta Zool. Lilloana, Tucuman 16: 13-21 (1958). 
Dibujos de los genitales masculinos de algunos satiridos neotropicales. (Lep. 
Rhop. Satyridae.) Acta Zool. Lilloana, Tucuman 16: 61-81 (1958). 

Butterfly migration in Argentina, Summer 1958. Entomol. 92: 28-41 (1959). 
Dos ninfalidos neotropicales nuevos. Neotropica 6: 30-32 (1960). 

Insectos Tucumanos perjudiciales. Rev. Indust. Agric., Est. Exper., Tucuman 
42: 3-144 (“1958,” 1960). [Also: Publ. Misc. No. 21, Instituto Miguel Lillo.] 
Migration of butterflies in northwestern Argentina, spring and summer 1959- 
1960. Entomol. 93: 237-240 (1960). 


Guia para el entomologo principante. Publ. Misc. No. 22, Inst. Miguel Lillo. 
168 p., Ghigs-2 7 pl: 

Migration of butterflies and moths in Argentina, spring and summer 1960-1961. 
Entomol. 95: 8-12 (1962). 

Satiridos Argentinos IV. Notas Adicionales (Lep. Rhop. Satyridae). Acta Zool. 
Lilloana, Tucuman 18: 11-17 (1962). 

Catalogo sinonimica de los ropaloceros Argentinos, excluyendo Hesperidae 
(Tercer suplemento.) Acta Zool. Lilloana, Tucuman 18: 19-30 (1962). 
Satiridos sudamericanos nuevos. Acta Zool. Lilloana, Tucuman 18: 105-109 

Dibujos de los genitales masculinos de algunos satiridos neotropicales. II. Acta 
Zool. Lilloana, Tucuman 18: 251-257 (1962). 

Migration of butterflies in northwestern Argentina, spring and summer 1961— 
1962. Entomol. 95: 237-239 (1962). 

Migration of butterflies in northwestern Argentina, late spring and summer 
1962-1963. Entomol. 96: 258-264 (1963). 

Dos satiridos neotropicales nuevos. Rev. Soc. Entomol. Argentina 26: 45—46 

. Tipos de Insecta (exceptuando Diptera), consevandos en el Instituto Miguel 

Lillo. Acta Zool. Lilloana, Tucuman 19: 297-334 (1964). 

. Tipos de Diptera conservados en el Instituto Miguel Lillo. Acta Zool. Lilloana, 

Tucuman 19: 391-406 (1964). 

Ropaloceras Argentinas. Observaciones varias. Acta Zool. Lilloana, Tucuman 
19: 487-492 (1964). 

Catalogo sinonimica de los ropaloceras Argentinas excluyendo Hesperididae. 
(Cuarto suplemento.) Acta Zool. Lilloana, Tucuman 19: 497-502 (1964). 

. Tipos de animales vertebrados conservados en el Instituto Miguel Lillo. Acta 

Zool. Lilloana, Tucuman 19: 509-510 (1964). 

Dibujos de los genitales masculinos de algunos satiridos neotropicales, III. 
Acta Zool. Lilloana, Tucuman 19: 511-517 (1964). 

Migration of butterflies in northwestern Argentina during the summer of 1964. 
Entomol. 97: 272-273 (1964). 

Una nueva Phyciodes de Peru. Acta Zool. Lilloana, Tucuman 20: 63-65 (1964). 
Euptychia weyrauchi, sp. nov. Acta Zool. Lilloana, Tucuman 20: 169-170 

Dibujos de los genitales masculinos de algunos satiridos neotropicales, IV. Acta 
Zool. Lilloana, Tucuman 20: 187-191 (1964). 

Insecta. Lepidoptera, fam. Nymphalidae et Fam. Heliconiidae. Tomo III del 
genera et species animalia Argentinorum. 474 p., 25 pl. (19 in color). Talleres 
Grafico Kraft, Buenos Aires (1964). 

Los ropaloceros de Cafayete (Salta.) Rev. Soc. Entomol. Argentina 28: 65-70. 

Notes on Argentine butterfly migration 1965-1966. Entomol. 100: 30-34 

Insecta. Lepidoptera, fam. Papilionidae et fam Satyridae. Tomo IV del genera — 
et species animalia Argentinorum. 412 p., 27 pls. (19 in color). Talleres 
Grafica Kraft, Buenos Aires (1967). 

Descripciones de Thecla. Acta Zool. Lilloana, Tucuman 21: 5-12 (1967). 
Una nueva Phyciodes de Ecuador Oriente. Acta Zool. Lilloana, Tucuman 21: 
13 —l5aGl9Gr)). 

. Ancyloxypha melanoura orientalis subsp. nov. Acta Zool. Lilloana, Tucuman 

Pe INS) (eIUSIS7/ })- 

. Tres satiridos nuevos. (Lep. Rhop. Satyridae) Acta Zool. Lilloana, Tucuman 

21: 20-24 (1967). 

VoLUME 28, NUMBER 4 343 

984. Nuevos nombres para generos de Lepidoptera. Acta Zool. Lilloana, Tucuman 
JI lao (1968). 

285. Dos especies nuevas de Audre. (Lep. Rhop. Riodinidae). Acta Zool. Lilloana, 
Tucuman 22: 137-142 (1968). 

286. Cuatro satiridos nuevas de Ecuador (Lep. Rhop. Satyridae). Acta Zool. Lillo- 
ana, Tucuman 22: 201-216 (1968). 

287. Fauna del noroeste Argentina. I. Las aves de Guayapa (La Rioja). Acta Zool. 
Lilloana, Tucuman 22: 211-220 (1968). 

288. Lista de los tipos de insects y otros invertebrados conservados el las colecciones 
de Instututo Miguel Lillo. (3a entrega.) Acta Zool. Lilloana, Tucuman 22: 
33/—352 (1968). 

289. Notes on butterfly migrations between April 1966 and May 1968. Entomol. 
102: 10-11 (1969). 

290. Datos para la estudio ontogenia de Lepidopteros Argentinos. Publ. Misc. No. 
31, Instituto Miguel Lillo, 142 p. (1969). 

291. Lista de los tipos de insectos y otros invertenbrados conservados en el Instituto 
Miguel Lillo. (4a entrega.) Acta Zool. Lilloana, Tucuman 26: 105-116 (1970). 

IP. Observations on migrations of Lepidoptera in northwestern Argentina, November 
1968—June 1970. Entomol. 

IP. Guia para el entomologi principiante. (Segunda Edicion). 

IP. Catalogo de los ropaloceros Argentinos. (450-500 p.). 


A very unusual copulation between two males and one female of Euphydryas 
chalcedona was observed on 13 June 1974 in Whitewater Canyon, San Bernardino 
Mountains, Riverside County, California. The three individuals were disturbed from 
a mating position at the top of a bush. Flight was attempted by the female who 
carried the two males with her. She was barely able to fly with this weight and the 
flight was on a downward trajectory until intercepted by my net. The three butter- 
flies were genitalically attached and appeared to be in copula. They remained so 
in the net and after being dispatched by pinching. All three individuals were very 
fresh, especially the female who had one hind wing incompletely opened. It is 
possible that this mating occurred before the female had flown. The time of day 
was 1300 PDT. 

A similar mating between two males and one female of Phyciodes phaon was re- 
ported by Perkins (1974, J. Lepid. Soc. 27: 291-294) who referred to it as a 
pleoheterosexual coupling. As far as I can ascertain, no other examples of a copula- 
tion between three or more individuals of Lepidoptera have been reported in the 

Joun H. Masters, 5211 Southern Avenue, South Gate, California 90280. 



Joun F. Emmet! anp THomMas C. EMMEL 
Department of Zoology, University of Florida, Gainesville, Florida 32611 

Ten years have passed since the publication of the last of a series of 
four papers (Emmel & Emmel, 1962a, 1962b, 1963a, 1963b) on the butter- 
fly associations and distributional factors affecting some 74 species of 
Rhopalocera in a montane ecosystem of the Sierra Nevada in California. 
With further study since our initially reported observations, a number 
of additional butterfly species have been recorded for this Donner Pass 
region and subsequent intensive invesetigations of hostplant relationships 
have been carried out by the authors and other California workers. A 
total of 83 butterfly species are now known for this four-square-mile area, 
making it the richest montane fauna of any reported temperate-zone area 
of comparable size (Emmel & Emmel, 1963b, p. 99). 

In the following report, species for which new hostplant information 
is known are listed by family name and the species number previously 
used in Emmel & Emmel (1962a). Some of the host identifications made 
in 1956 and 1960 in the Donner Pass region have been changed from the 
original botanical names given to the the authors, and listed in Emmel & 
Emmel (1962a), due to subsequent submission of voucher material to 
other botanical specialists. The butterfly species new to the previously- 
recorded fauna are also listed (with an asterisk) at the end of each family 
section, with numbers subsequent to those for species recorded originally 
for the family. 


1. Papilio zelicaon zelicaon Lucas. Previously recorded on “Cymopterus tere- 
binthinus,” this foodplant is now known as Pteryxia terebinthina (Hook.) C. & R. 
var. californica Math. 

2. Papilio indra indra Reakirt. The change in host identification is identical to 
that for P. zelicaon. 

4. Papilio eurymedon Lucas. The foodplant in this locality is now strongly 
suspected to be Ceanothus cordulatus Kell., from observations of females hovering 
around this particular species. 

5. Parnassius clodius baldur Edwards. Females have been observed ovipositing 

1 Research Associate, Natural History Museum of Los Angeles County. Mailing address: 1117 
Oth Street, Santa Monica, California 90403. 

VoLUME 28, NUMBER 4 345 

on or near Sedum obtusatum Gray (Emmel & Emmel, 1962a); however, this Sedum 
is only one of several oviposition substrates and is not a foodplant, as the normal 
host for the larvae appears to be Dicentra uniflora Kell. (Fumariaceae) (J. F. Emmel, 
unpubl.), which is abundant right after snow melt on the slopes where baldur flies 
later in the summer (when the Dicentra plants are dried and shriveled, e.g. on 18 
July 1970). 


3. Pieris occidentalis Reakirt. All of our material previously called P. protodice 
is now referable to P. occidentalis, a sibling species as defined by Chang (1963). 

5. Euchloe hyantis hyantis Edwards. The foodplant on the lower slopes of Mt. 
Judah is Streptanthus tortuosus Kell. var. orbiculatus (Greene) Hall (Cruciferae ). 
Several females were observed (18 July 1969) to oviposit on the buds of this mustard. 

6. Anthocharis sara stella Edwards. Name changed from A. s. julia Edwards. 
Two local food plants for this butterfly are Arabis platysperma Gray (females ovi- 
positing 21 June 1970, and ova plus larvae found on 11 July 1971) and Arabis lyallii 
Wats. (ova on 21 June 1970). 

*10. Pieris protodice Boisduval & LeConte. Shapiro (1974) took this species 
from 27 June to 28 September 1973 in the Donner Pass area. 

“11. Pieris beckerii Edward. This species was recorded by Shapiro (1974) from 
25 July to 24 August 1973 in the Donner Pass area. 


2. Cercyonis oetus oetus (Boisduval). Name change from C. sthenele oetus (see 
Emmel, 1969). 

*3. O0ceneis ivallda (Mead). The presence of this species was first called to 
our attention by Noel La Due (in litt., 7 August 1963). He found it on the north 
slope of Mt. Judah on 16 July 1963 in fair numbers. One of us (JFE) in company 
with Oakley Shields took ivallda on 15 June 1969, 18 July 1969 and 11 July 1971. 
(The same areas had been checked in 1970, with no adults seen.) Our previous 
Donner Pass collections were made in 1956 and 1960. Thus the Oeneis ivallda pop- 
ulations in the Donner Pass region seem to be synchronized to fly only in odd- 
numbered years (a two-year cycle is well known for Oeneis nevadensis Felder & 
Felder and O. jutta reducta McDunnough in California and Colorado). The suspected 
foodplant is Carex spectabilis Dewey, with which females have been observed to 
be closely associated. This locality, at 7200 ft. elevation, is the lowest altitudinal 
record known for Oeneis iwallda. 


9. Phyciodes campestris montana Behr. An egg mass of 118 eggs was found 
on the underside of a leaf of Aster occidentalis (Nutt.) T. & G. in a wet meadow 
near Lake Mary on 21 August 1971. This Aster species is probably the same species 
we referred to (Emmel & Emmel, 1962a) as Aster integrifolius Nutt. 

11. Polygonia zephyrus Edwards. Correct name for foodplant is Ribes cereum 
Dougl. rather than R. viscosissimum Pursh. 

18. Cynthia annabella Field. Name change from Vanessa carye Hiibner. Host- 
plant here is Sidalcea glaucescens Greene (Malvaceae ). 

*21. Speyeria egleis egleis (Behr). This species is about as abundant as the 
phenotypically very similar Speyeria mormonia arge (Strecker) throughout the Pass 
area. Observations during July 1969 showed that S. egleis prefers dry meadows and 
montane slopes while S. mormonia prefers forest edges. 

*22. Adelpha bredowii californica (Butler). On 21 June 1970, three females 


were collected on the slope between Donner Pass and Mt. Judah at elevations be- 
tween 7200 and 7500 feet (perhaps a new altitude record for this subspecies). One 
of these females was observed to oviposit on a leaf tip of Quercus vaccinifolia Kell. 
Adelpha has not been observed previously in the Pass region, and this 1970 record 
may represent merely an occasional high-altitude invasion by a predominantly low- 
land species. 


3. Satyrium saepium (Boisduval). This hairstreak feeds on a_small-leafed 
Ceanothus (unidentified), previously erroneously identified for us as C. velutinus 
Dougl. ex Hook. 

4. Satyrium behrii (Edwards). The foodplant in the Donner Pass area is Purshia 
tridentata (Pursh) D.C., which grows on the ridge and upper slopes of Mt. Judah. 

5. Satyrium fuliginosum (Edwards). On 15 June 1969, seven mature larvae 
were collected in debris and soil at the base of plants of Lupinus arbustus Douglas 
on the summit of Mt. Judah. 

8. Mitoura nelsoni nelsoni (Boisduval). On 18 July 1970, a worn male was 
taken in association with Juniperus occidentalis Hook. on the north slope of Mt. Judah 
(at lower elevations this butterfly is commonly associated with Calocedrus decurrens 
Torr., which is absent from Donner Pass). We suspect this Juniperus to be the 
food plant of M. nelsoni at Donner Pass. 

11. Callophrys lemberti Tilden. Name change from C. dumetorum perplexa 
Barnes & Benjamin. Suspected foodplant is Eriogonum marifolium Torrey & Gray, 
with which the adults are very closely associated on the slopes of Mt. Judah. 

12. Lycaena arota virginiensis Edwards. The previously misidentified host spe- 
cies is actually Ribes roezlii Regel, not R. montigenum McClat. 

13. Lycaena editha Mead. Two females were observed to oviposit in ground 
litter near Rumex acetosella L., near Lake Mary on 21 August 1971. This introduced 
dock is common throughout the Donner Pass meadows. 

14. Lycaena nivalis Boisduval. The foodplant is Polygonum douglasii Greene in 
other Pacific montane areas (Newcomer 1911, 1964); since this plant grows in the 
Donner Pass area, it is very probably the foodplant here. 

15. Lycaena cupreus (Edwards). The host plant is Rumex acetosella L. in the 
Lodge Meadow and Lake Mary areas. Previous recordings of unreared larvae on 
Calyptridium are almost certainly not this copper, but more probably larvae of 
Strymon melinus Hubner. 

18. Lycaeides argyrognomon anna (Edwards). On 18 July 1969 a female was 
observed to oviposit on a Lupinus species on the north slope of Mt. Judah; the 
plant lacked flowers or fruit and could not be identified as to species. 

2. Plebejus shasta (Edwards). On 15 June 1969 six mature larvae were found 
on flowers of and at the bases of plants of Astragalus whitneyi Gray on the north 
slope of Mt. Judah. 

23. Plebejus lupini Boisduval. Goodpasture (1973) has raised this taxon to 
specific level (name change from that used in Emmel & Emmel (1962a)). Adults 
are closely associated with Eriogonum umbellatum Torrey, the suspected foodplant 
at Donner Pass. 

24. Agriades glandon podarce Felder & Felder. The foodplant at Donner Pass 
is apparently Dodecatheon speices (Primulaceae ), based on close association of adults 
with these plants. 

25. Glaucopsyche lygdamus columbia Skinner. Name change from G. l. behvrii 
(Edwards) (after Langston, 1969). 

27. Philotes battoides intermedia Barnes & McDunnough. The adults are closely 
associated with Eriogonum umbellatum Torrey, the probable foodplant. 

VoLUME 28, NUMBER 4 347 

*99. Incisalia fotis windi Clench. On 10 July 1971 three first-instar and six 
fourth-instar larvae were found feeding on flowers and leaves of Sedum obtusatum 
Gray near Lake Mary. No adults had been previously recorded for the area. 

*30. Lycaena mariposa Reakirt. On 17 July 1963 Noel La Due took three adults 
on a north-facing slope near Lake Mary. 

*31. Apodemia mormo (Felder & Felder). On 17 August 1973, Shapiro took 
this species at Donner Pass. Details on its habitat will be published by him in a 
future paper. We had previously noted its suspected Sierran foodplant, Eriogonum 
wrightii Torr. ex Benth., to be abundant on the granite rock slopes at Donner Pass. 


1. Thorybes nevada Scudder. This skipper is closely associated with a Trifolium 
(Leguminosae) species, the suspected foodplant. 

4. Erynnis propertius (Scudder & Burgess). Determination change from E. 
juvenalis (Fabricius) (C. D. MacNeill, in litt.). The foodplant at Donner Pass is 
strongly suspected to be Quercus vaccinifolia Kell. 

5. Erynnis persius (Scudder) complex. Determination change from E. afranius 
(Lintner) (Burns 1964). 

*11. Ochlodes sylvanoides (Boisduval). Shapiro has taken this species in the 
Donner Pass region (17 August-5 October 1973) and will publish detailed data 
on its ecological associations in a future paper. 


1. Parnassius phoebus behrii Edwards. Specimens of this species were seen on 
Mt. Anderson and Tinker Knob in 1960, south of the Donner Pass area; the ridge 
connecting these peaks also extends to Mt. Lincoln, just inside the study area 
delineated in our 1962a paper. Because Mt. Lincoln is similar in habitat and flora 
to these peaks south of it, it is probable that this Parnassius will be eventually be 
taken there, also. 


Burns, J. M. 1964. The skippers of the genus Hesperia in western North America 
(Lepidoptera: Hesperiidae). Univ. Calif. Publ. Entomol. 35: 1-230. 

CuHanec, V. C. S. 1963. Quantitative analysis of certain wing and genitalia char- 
acters of Pieris in western North America. J. Res. Lepid. 2: 97-125. 

EMMEL, T. C. & J. F. Emmet. 1962a. Ecological studies of Rhopalocera in a 
High Sierran community—Donner Pass, California. I. Butterfly associations and 
distributional factors. J. Lepid. Soc. 16: 23-44. 

1962b. Note regarding habitats and Rhopalocera of Donner Pass, Cali- 

fornia. J. Lepid. Soc. 16: 136. 

1963a. Ecological studies of Rhopalocera in a High Sierran community— 

Donner Pass, California. II. Meteorologic influence on flight activity. J. Lepid. 

Soest: 7-20. 

1963b. Composition and relative abundance in a temperate zone butter- 
fly fauna. J. Res. Lepid. 1: 97-108. 

GooppastTuRE, C. 1973. A new subspecies of Plebejus acmon (Lepidoptera: 
Lycaenidae). Pan-Pac. Entomol. 49: 149-159. 

Laneston, R. L. 1969. A review of Glaucopsyche, the silvery blues, in California 
(Lycaenidae). J. Lepid. Soc. 23: 149-154. 

Newcomer, E. J. 1911. The life histories of two lycaenid butterflies. Canad. 
Entomol. 43: 83-88. 


—. (“1963”) 1964. The synonymy, variability and biology of Lycaena nivalis. 
J. Res. Lepid. 2; 271-280. 

SHaprro, A. M. 1974. Ecological and behavioral aspects of coexistence in six 
Crucifer-feeding Pierid butterflies in the central Sierra Nevada. Amer. Mid. 
Nat. (in press). 


During the summer of 1972 the second author was collecting butterflies in northern 
British Columbia. Among the many extensions of butterfly distribution noted, three 
were of uncommon interest. These were all collected at one locality and in one 
day: Mt. Hoadley, near New Aiyansh, British Columbia, 55° 128° SW, 19 July 1972. 
One female of Parnassius eversmanni Ménétriés was collected. Males were observed 
but not collected. This represents a new species for British Columbia. It also rep- 
resents a 430-mile southern extension of the species’ known range in Mt. McKinley 
National Park and Eagle Summit, Alaska. It is not clear from the single female if 
this population is referable to P. e. thor Hy. Edwards. 

Boloria epithore chermocki Perkins and Perkins (two males, two females) and 
Erebia vidleri Elwes (two females) were also collected. Evrebia vidleri was not 
previously known north of Vancouver, British Columbia on the coast and Lillooet, 
British Columbia in the interior. Except for doubtful records from central Alaska 
(which have not been confirmed by the Alaska Lepidoptera Survey) and doubtful 
records for Smithers and Chilcotin, British Columbia, B. epithore was known posi- 
tively only south of Lillooet, British Columbia. Thus the known range for B. epithore 
and E. vidleri has been extended 400 miles to the north. 

Parnassius eversmanni was taken above timberline (6400 ft.) and replaced P. 
phoebus Fabricius which was just at timberline (5000 ft.). Boloria epithore and 
Erebia vidleri were taken at 5500 ft. elevation. At the lower elevations, Plebejus 
saepiolus (Boisduval), Pieris napi (Linnaeus), Speyeria mormonia (Boisduval), S. 
hydaspe (Boisduval), and Papilio zelicaon Lucas were also taken. This is a common 
species association for Boloria epithore and Erebia vidleri at more southern coastal 

In addition to the above records, one female of Boloria epithore was recorded at 
the following, more inland, locality: Hudson Bay Mountain, Ski Hill, near Smithers, 
British Columbia, 26 July 1972. The other two species were not seen here. This 
locality is near that for a doubtfully accurate record of Parnassius eversmanni that 
has been overlooked or ignored by other authors (Jones 1951, Entomol. Soc. Brit. 
Columbia, Occ. Pap. 1, 148 p.). Gunder (1932, Pan-Pac. Entomol. 8: 123-127) 
recorded Parnassius eversmanni as follows: Babine Range above Smithers, British 
Columbia, 20 July 1931, J. F. May, one female. The Smithers record for B. epithore 
(Perkins & Meyer 1973, Bull. Allyn Mus. Entomol. 11: 1-23) is the same as Par- 
nassius eversmanni. Since the species Melitaea mayi Gunder from the same locality 
is of doubtful existence anywhere in North America, it was assumed that the Par- 
nassius eversmanni and Boloria epithore were similarly mislabeled. 

Jon H. SHeparD AND Sicrip M. SHEeparD, Rural Route #2, Nelson, British Co- 
lumbia, Canada. 

VoLuME 28, NUMBER 4 349 


James E.. Lioyp 
Department of Entomology, University of Florida, Gainesville, Florida 32611 

Males of Psilogramma menephron (Cramer) and Psilogramma jordana 
Bethune-Baker produce sounds by rasping scales on the dorsal surfaces 
of the genitalic valves against needle like spines that are located on the 
posterior edge of the eighth tergite. There are no significant differences 
between the stridulatory structures of the 2 species. Temporal charac- 
teristics of the sound of P. jordana have been described by Robinson & 
Robinson (1972), and I recorded the sounds of P. menephron at the 
Bishop Museum Field Station (Wau Ecology Institute) at Wau, New 
Guinea.! Its sound differs from that reported for P. jordana. The con- 
clusions of Robinson & Robinson (1972) regarding the frequency output 
of the stridulatory mechanism are incorrect. 

The moths were taken at night at an incandescent bulb, and when 
grasped and manipulated in the hand, they produced sibilant tss_ tss 
sounds. These sounds were emitted in groups that were irregular in 
duration (from less than 1 to more than 4 seconds) and rhythm. 

Analysis of the recorded sounds? reveals the following: each tss sound 
is composed of a variable number of acoustical units (pulses), and there 
is no structure within a pulse that would suggest the actual nature of 
the spine-scale stridulatory mechanism (Fig. 1A). The sound spectrum 
is continuous from 1 to about 14 kilohertz (Fig. 1B) and within this 
range there are no especially dominant frequencies. The time charac- 
teristics of the pulses and periods of portions of 3 pulse groups are given 
in Table 1. Pulse frequencies were 11.1-12.5 Hz (26.5°). 


The sounds of the 2 species are similar though not identical. The pulses 
of P. menephron were emitted in groups, but no grouping is apparent 
in the pulse train figured by Robinson & Robinson (1972) for P. jordana. 
The pulse recurrence frequency of P. menephron (ca 12 Hz at 26.5°) is 
about 2X that of P. jordana (temperature unknown). Pulse length in P. 
jordana is about 2X that of P. menephron (0.14 versus 0.07 sec). 

1 This research was performed during the 1969 Alpha Helix Expedition to New Guinea; the 
program was supported by the National Science Foundation under grant GB 8400 to the Scripps 
Institution of Oceanography. 

2 Recordings were made with a Uher 4000 Report-L tape recorder at 7.5 ips, and an Electro- 
Voice 655C dynamic microphone. Analysis was made with a Sona-Graph with an analyzing band 
width of 300 Hz. 


i ; t i 

Fig. 1. Moth sounds: (A) audiospectrogram showing a long sequence of pulses 
(vertical axis = frequency in kilohertz; horizontal axis = time in seconds; (B) audio- 
spectrogram with frequency and time axes doubled, showing the broad carrier fre- 
quency spectrum; (C) oscillogram showing pulse of sound with its beam deflection 
frequency of several thousand (estimated 10,000) per second. Recording temperature 
for all sounds figured was 26.5°C. 

It is not possible to determine the actual mechanics of sound produc- 
tion at the level of spine-scale impact from simple tape recordings, as 
attempted by Robinson & Robinson (1972). They counted oscilloscopic 
beam deflections, compared this with counts of spines and scales, and 
suggested that each beam deflection was a spine-scale impact (they 
estimated an impact frequency of 1,430 Hz). The audiospectrogram 
(Fig. 1B) shows that the spectrum is continuous from 1 to at least 14 

VoLUME 28, NUMBER 4 Sie 

TaBLE 1. Pulse characteristics of portions of three pulse groups. 

Pulse Length (sec) Pulse Period (sec) 
Grou ———E———ESS-- Nae PES AES cee 
No. x Range s.d. n. x. Range s.d. n. 
il 0.07 0.06-—0.09 0.01 19 0.09 0.08—0.13 0.01 19 
Z 0.07 0.03—0.08 0.01 TS} 0.08 0.07—0.12 0.01 22 

3 0.07 0.03-0.11 0.02 23 0.09 0.06—0.12 0.12 22 

KHz, and no spine-scale impact frequency is evident amid the myriad of 
carrier frequencies. (1) The oscilloscope beam deflection is the result 
of averaging hundreds of frequencies at many different energy levels. 
(2) Even in the simple (by comparison) file and scraper stridulation 
of Tettigoniidae, a 1 to 1 relation does not exist between unit impact 
and sound output: the acoustical output of a single file-tooth impact is 
a complex wave of several cycles (Sugo, 1966). (3) Actually, the oscillo- 
gram given for P. jordana does not appear to be completely resolved into 
individual beam deflections. By using a fine, low-intensity beam, fast 
film, and a sweep speed of 100 cm/sec I was able to resolve the sound 
of P. menephron to an estimated 10,000 beam deflections/second (Fig. 
1C),® a figure of no real meaning or descriptive significance when 
compared with the acoustical parameters that were determined audio- 


I acknowledge the many kindnesses of and assistance received from 
colleagues on the Expedition and friends and associates in the T.P.N.G. 
Among these were J. and E. Buck, M. and J. Sedlacek, and J. Wormersley 
and his staff of the Lae Botanical Garden. I thank T. J. Walker, J. J. 
Whitesell and S. M. Ulagaraj for technical advice and assistance. T. J. 
Walker read the manuscript. This research was performed during the 
tenure of N.S.F. grant GB 7407, Florida Agricultural Experiment Station 
Journal Series no. 5362. 

3 Tektronix oscilloscope 564, 3A72 amplifier, and 2B67 time base unit; Tektronix C30 camera. 


Rosinson, G. S. & H. S. Roprnson. 1972. Genital stridulation in male Psilogramma 
jordana Bethune-Baker (Lepidoptera, Sphingidae). Entomol. Rec. & J. Var. 
84: 213-215. 

Suca, N. 1966. Ultrasonic production and its reception in some neotropical Tet- 
tigoniidae. J. Insect Physiol. 12: 1039-1050. 



J. W. TitpEN 
125 Cedar Lane, San Jose, California 95127 

Tilden (1969) considered the name pulchella Boisduval a synonym 
of tharos Drury 1773. Some question was raised about this action. It 
was pointed out that there is in the United States National Museum, a 
specimen that is labelled as the type of Melitaea pulchella Boisduyal. 

W. D. Field kindly examined this specimen and stated that he con- 
sidered it a specimen of Phyciodes campestris campestris (Behr) but 
agreed with my opinion that Boisduval had not described this insect in 
his original statement concerning pulchella. He expressed the belief that 
pulchella was a replacement name, since, as pointed out by Tilden (1969), 
Boisduval had merely cited Drury’s figure as representing his Melitaea 

In July 1973 I was able to examine the type of M. pulchella. It is 
indeed a specimen of Phyciodes campestris, without locality or date 
labels. It is thus not possible to be sure when this specimen was selected 
by Boisduval. It may have been at the time of the original citation, or 

Boisduval’s statement that Melitaea pulchella (which he considered 
to be represented by figs. 5 & 6 on Plate 1 of Drury’s Illust. Nat. Hist. ) 
should not be confused with Papilio tharos Cramer, indicated that he 
gave priority to tharos Cramer and thus intended pulchella as a replace- 
ment name for Papilio tharos Drury. 

There is no description of any insect, either here or in later references 
to pulchella by Boisduval. 

It makes no difference what insect is labelled as the type of pulchella, 
since this insect so labelled is not described and so is without status. By 
Boisduval’s own statements, pulchella is a replacement name for Papilio 
tharos Drury, not Papilio tharos Cramer. 

Cramer (Tom. II p. 12, & Plate CLXIX, figs. E, F) figures Papilio 
tharos and refers to Drury, Tom. I, pl. 12, figs. 5, 6. Papilio tharos Drury 
dates to 1773, P. tharos Cramer to 1777. 

On the basis of Boisduval’s statements, Melitaea pulchella must be 
considered a synonym of Papilio tharos Drury, and cannot replace 
Phyciodes campestris (Behr) 1863 even though the “type” of pulchella 
is a specimen of campestris. 

VoLUME 28, NUMBER 4 55) 


BotspuvAL, J. 1852. Ann. Soc. Entomol. France (2)10(2): 306, no. 4. 

. 1869. Ann. Soc. Entomol. Belg. 12: 20, no. 50. 

Crammumeeely (9 (1777). Uit. Kap:, Tom II, p. 12; pl. CILXIX, figs: E, F. 

Devewew miro. Iiust. Nat. Hist., pl: 21, figs. 5, 6: 

Titpen, J. W. 1969 (1970). Concerning the names and status of certain North 
American members of the genus Phyciodes. J. Res. Lepid. 8(4): 94-98. 


Harris (1972, Butterflies of Georgia, Univ. Oklahoma Press) reports that John C. 
Symmes found and reared Satyrium kingi (Klots & Clench) on Flame Azalea ( Rho- 
dodendron calendulaceum ) in the Atlanta, Georgia area; but that H. L. King collected 
kingi at the type locality (Savannah, Georgia), where he saw females ovipositing on 
a small plant not related to azalea. Moreover, King noted that he found no native 
azalea plants in the area around where he collected his specimens. These facts, of 
course, suggest that kingi has more than one foodplant. More recently Gatrelle 
(1974, J. Lepid. Soc. 28: 33-37) has raised the question of the relationship between 
possible subspecifically distinct populations of kingi and differences in the choice of 
foodplant in these different populations. More specifically, the inference might be 
made that the northern (inland or upland) population not only represents a sub- 
species distinct from the lowland (or coastal) population, but that the northern pop- 
ulation may feed on a different foodplant from the lowland population. 

I wish to report a second foodplant for the northern population of kingi, horse 
sugar tree, Symplocos tinctoria (L.). On 10 May 1966, on a ridge near the Chatta- 
hoochee River just north of Atlanta, Georgia, I found three larvae that were un- 
familiar to me on a single bushy plant. The three larvae, along with an ample supply 
of the foodplant, were collected; and the larvae were reared at my home in Atlanta. 
On 17 May 1966 the first larva pupated and the other two pupated several days 
later. The first adult emerged on 28 May 1966 and the other two emerged several 
days later. Upon identifying the specimens as Satyrium kingi, I pressed a branch 
of the foodplant (which was still quite fresh even 18 days after it had been collected). 
The foodplant was later identified as horse sugar tree by Dr. Robert Godfrey, De- 
partment of Botany, Florida State University. The larvae I reared fit the general 
description given by Harris (loc. cit.), and were similar in appearance to a single 
larva of Satyrium liparops (Boisduval & Le Conte) which I collected almost a year 
later (2 April 1967) on wild cherry (Prunus sp.) less than 300 meters from the spot 
where the kingi larvae were found. The liparops larva pupated on 6 April 1967 and 
the adult emerged on 16 April 1967. 

Single adult specimens of kingi were collected in the same general area of upland 
hardwoods on 3 June 1966 and 9 June 1967. Other members of the family Lycaenidae 
that I collected at the same location in 1966 and 1967 included Chrysophanus titus 
mopsus (Hubner) on 9 June 1967; Satyrium edwardsii (Grote & Robinson) on 9 
June 1967; Strymon melinus (Hiibner) on 9 June 1967; Satyrium calanus falacer 
(Godart) on 3 June 1966; Calycopis cecrops (Fabricius) on 17 April 1967; Atlides 
halesus (Cramer) on 13 March 1967; and Callophrys augustinus croesides (Scudder ) 
on 13 March 1967. 

J. C. Fioyp, 5106 Arrowhead Drive, Baytown, Texas 77520. 



3026 Bapaume Avenue, Norfolk, Virginia 23509 

A year’s collecting in the Lower Florida Keys during 1972-1973 turned 
up three species of butterflies not previously recorded for the United 
States and several other uncommon and unusual captures. It is difficult 
to estimate on the basis of the present records how extensively the new 
species have established themselves, but at least one has a well estab- 
lished colony. 

Electrostrymon angelia angelia (Hewitson ) 
(Fig. 1) 

This species was first captured in Key West on 6 April 1973 and was 
taken continuously each month until my departure in November 1973. 
E. angelia was found in and around a tropical hardwood area and was 
attracted to the blossoms of Brazilian Pepper Schinus terebinthefolius 
(Raddi) and Seagrape Coccoloba uwifera (L.). When these blossoms 
were no longer available, the butterflies were found perched on leaves at 
the edge of the wooded area or in open areas within the trees, and were 
almost always perched in areas of shade or broken sunlight rather than 
in direct sunlight. A preference for shaded areas was especially notice- 
able in the summer months, whereas in April and again in October-— 
November perches were more likely to be in areas of scattered sunlight. 

E. angelia was not common and seldom were more than 3-5 specimens 
captured at one location. However, during the end of April as many 
as 20-25 specimens were seen flying about in clearings and open areas 
on clear, hot afternoons. This hairstreak did not seem to prefer any 
particular height for perching, rather the nature of the foliage and the 
amount of sunlight seemed to determine the perch. 

Harry Clench of the Carnegie Museum has determined that the Key 
West population belongs to the nominate populations found on Cuba 
and not to E. angelia dowi (Clench) which occurs in the Bahamas. — 
Apparently, the colony on Key West is well established and should 
remain barring destruction of the area. Specimens are being deposited 
in the collections of the Carnegie Museum, Pittsburgh, Pennsylvania, 
and the Allyn Museum, Sarasota, Florida. 


VoLUME 28, NUMBER 4 apo 

Fig. 1. Electrostrymon angelia angelia (Hewitson), ¢, upper (left) and under 
(right) surfaces (collected at Key West, Munroe Co., Florida, 30 May 1973, R. A. 
Anderson leg.) 2.7. Allyn Museum photo nos. 110773-15/16. 

Strymon limenia (Hewitson) 
(Fig. 2) 

Two males and one female of this species were captured on 23 
December 1972 on Big Pine Key on the flowers of Spanish Needles 
(Bidens pilosa L.). Had I not had previous experience with S. limenia, 
I’m sure I would have overlooked it due to the similarity in pattern 
between this species and the more common S. columella modesta (May- 
nard). A single male S. limenia was also captured in Key West on 23 
May 1973 when resting near blossoms of a Brazilian Pepper. Subsequent 

Fig. 2. Strymon limenia (Hewitson), 6, upper (left) and under (right) surfaces 
(collected at Key West, Munroe Co., Florida, 23 May 1973, R. A. Anderson leg.). 
2.7. Allyn Museum photo nos. 110773-17/18. 


collecting in Key West and on Big Pine Key did not produce additional 
specimens. Steve Roman of Orlando, Florida has reported (in litt.) that 
he found a male S. limenia dated 3 April 1971 in his series of S. columella 
from Big Pine Key. 

The known records for this species cover a period of two years and 
two Keys which are thirty-five miles apart. Perhaps this hairstreak is 
established on the Keys between the known locations as well as on other 
Keys toward the mainland, and has heretofore escaped detection due to 
its similarity to S. columella. One male and one female have been de- 
posited in the collection of the Allyn Museum. 

Anartia lytrea (Godart) 

A fresh male of this species was captured on 22 February 1973 in 
Key West. Harry Clench has mentioned (pers. comm.) that he has seen 
another specimen of A. lytrea from Big Pine Key captured in 1972. These 
two records, thirty-five miles apart, suggest the possibility of an estab- 
lished colony in the Lower Keys. The specimen from Key West was 
flying in an open wooded area and conveniently landed on the ground 
where it was captured. This particular locality was frequently visited 
but no additional specimens were seen. Lee Miller of the Allyn Museum 
has indicated (pers. comm.) that the specimen from Key West does not 
match the description of A. lytrea chrysopelea (Hubner) from Cuba and 
therefore the proper subspecific determination has not been made at 
this time. My specimen has been deposited in the collection of the Allyn 

Chlorostrymon maesites maesites (Herrich-Schaffer ) 

Twenty-seven specimens of this rare hairstreak were taken in Key West 
during the months of May through September 1973. Although there are 
captures for each month during this time, most of the specimens were 
captured from the last week in May to the middle of June. This peak 
in numbers occurred when the Brazilian Pepper and Guamachil Apes- 
earring (Pithcellobium dulce Benth.) were in bloom. Blossoms of 
Pithcellobium were especially attractive. Although some C. maesites 
were captured while visiting the blossoms of these two trees, most were 
found perched on leaves of other nearby trees, and always on perches 
in direct sunlight. No more than half of the individuals seen were cap- 
tured because they constantly changed perches, their flight being rapid 
and difficult to follow, and frequently their perches were out of range 
of my long-handled (12 ft.) net. However, some specimens were taken 

VoLuME 28, NuMBER 4 357 

Fig. 3. Anartia lytrea (Godart), ¢, upper (left) and under (right) surfaces 
(collected at Key West, Munroe Co., Florida, 22 February 1973, R. A. Anderson 
leg.). 1.5. Allyn Museum photo nos. 110773-13/14. 

from perches as low as 3-6 ft. above the ground. Its small size and green 
underside often made this hairstreak extremely difficult to see when 
resting on foliage. Despite the moderate number of C. maesites taken, 
it was not common on Key West and the individuals seen and captured 
were the result of frequent visits to the known colonies for a period of 
several months. 

Marpesia eleuchia (Hubner) 

A male of this Antillean Dagger Wing was captured on 14 October 
1973 on Sugarloaf Key. It was attracted to the blossoms of a Brazilian 
Pepper and was flying with M. petreus (Cramer) which was common 
at the time. The specimen was in good condition and did not look as 
though it was a visitor from outside the Keys. Subsequent visits to the 
area during the rest of October did not locate additional specimens. The 
specimen has been deposited in the collection of the Allyn Museum. 

Eurema boisduvaliana (Felder) 

One fresh female was taken on 20 September 1973 in an open wooded 
area in Key West. The capture was made in a frequently collected area 
and was the only example of this species seen. The specimen has been 
deposited in the collection of the Allyn Museum. 

Erynnis zarucco zarucco (Lucas) 

Approximately twenty percent of the specimens seen from the Key 
West area have a white fringe on the hindwing, which is characteristic 


of E. zarucco funeralis (Scudder and Burgess). The white fringe is not 
as extensive as in typical E. z. funeralis but is intermediate between it 
and typical E. z. zarucco. Most of the specimens with significant white 
fringes on the hindwing were females. Interestingly, Kimball (1965, 
Lepidoptera of Florida) records four male funeralis-like E. zarucco from 
northern Florida. 


I wish to thank Dr. Lee Miller of the Allyn Museum for making the 
photographs accompanying this article, and Harry Clench of the Car- 
negie Museum for his subspecific determination of E. angelia. My ap- 
preciation also goes to Stan S. Nicolay and Dr. J. Bolling Sullivan for 
their suggestions and critical reviews of the manuscript. 


The first recorded hybrid between Colias eurytheme (Boisduval) and Colias har- 
fordii (H. Edwards), a perfect male, was captured by the author near Cachuma 
Creek, San Rafael Mountains, Santa Barbara County, California. The date of capture 
was 1 May 1970. This locality is approximately two miles south of Cachuma Saddle 
Ranger Station, and five miles southeast of Figueroa Mountain. Adults of both 
eurytheme and harfordii have been observed flying in the San Rafael Mountains, 
and larvae of both species have been found on Astragalus antisellii (Gray) in Oso 
Canyon. The specimen has been placed in the Peabody Museum of Natural History, 
at Yale University, New Haven, Connecticut. 

RicHArp C. PriesraF, 5631 Cielo Avenue, Goleta, California 93017. 

VoLUME 28, NuMBER 4 359 


126 Wells Road, Hanahan, South Carolina 29405 
114 Monica Boulevard, Savannah, Georgia 31400 

The appalachian eyed brown, Lethe appalachia (R. L. Chermock), 
was first described as a subspecies of Lethe eurydice (Johansson) rang- 
ing from the mountains of West Virginia southward through the Ap- 
palachian Mountains and into northern Florida. It was also mentioned 
as occurring in the coastal swamps of Virginia and South Carolina 
(Chermock, 1947). 

Cardé et al. (1970) recognized L. appalachia as a distinct species that 
is broadly sympatric with L. eurydice but occurring mainly in swamp 
forests, shrub swamps and forest-edge ecotones, while eurydice occurs 
in open marshes and sedge meadows. Carde et al. (1970) gave the range 
of appalachia as Maine to northern Florida and westward to South 
Dakota and Alabama. 

The type locality of L. appalachia appalachia is Brevard, Transylvania 
County, North Carolina in the area of Connestee Falls. Nominate ap- 
palachia until recently had not been recorded from very many localities 
in the southern states. The known southern limit of the species was ex- 
tended in 1972 by the discovery of a colony in a swampy forest in west 
central Florida, two miles south of Zephyrhills, Pasco County (Brown, 
1973). The western range of appalachia appalachia was extended when 
it was found in Tishomingo County, Mississippi, by Mr. C. T. Bryson in 
May 1971. 

The northern populations of L. appalachia ranging from Massachusetts 
and Maryland westward to Wisconsin and Illinois were found to be 
sufficiently distinct from southern nominate appalachia populations to 
warrant a subspecific name. We name this new subspecies for Mr. Irwin 
Leeuw of Cary, Illinois, who first drew it to our attention through speci- 
mens that he collected in Michigan. 

Lethe appalachia leeuwi (Gatrelle and Arbogast), new subspecies 
(Figs. 5-14) 

Male: Forewing radius: 21-26 mm, mean 24.2 mm in type series. Dorsal surface: 
ground color of both primaries and secondaries grayish brown as in nominate sub- 


4 8 

Figs. 1-4. Lethe appalachia appalachia (R. L. Chermock): 1 & 3, male, Table 
Rock State Park, Pickens County, South Carolina, 3 July 1972; 2 & 4, female, Mc- 
Clellanville, Charleston County, South Carolina, 30 May 1970. 

Figs. 5-8. Lethe appalachia leeuwi (Gatrelle & Arbogast), new subspecies: 5 & 
7, holotype, male, Wakelee, Cass County, Michigan, 24 July 1972; 6 & 8, allotype 
female, Wakelee, Cass County, Michigan, 4 July 1957. 

VoLUME 28, NUMBER 4 361 

species (Figs. 1 & 5), but lighter and with more contrast between the various shades. 
Apical and marginal areas dark; narrow dark bar at end of cell and dark postmedian 
band of irregular width extending from Rl to CU1 and CU2. Area between this 
band and the row of ocelli light. Basal and discal areas an intermediate shade. 
Contrast between discal and limbal areas of secondaries more pronounced than in 
nominate appalachia and light rings surrounding ocelli usually more conspicuous. 
Ventral surface: ground color lighter and much less uniform than in nominate sub- 
species (Figs. 3 & 7) and lacking purplish cast. On both primaries and secondaries 
a broad band of light brown tinged with white extends from near costal margin to 
second anal vein. This band contrasts markedly with darker basal and discal areas. 
It is bordered proximally on both wings by the postmedian line, and on the primaries 
bordered distally by the row of ocelli. On the secondaries it surrounds the first 
ocellus and is bordered distally by the remaining ocelli. Female: Forewing radius: 
26-27 mm, mean 26.5 mm. As in male but lighter and with even more contrast 
between the various shades. Light area on dorsal surface of primaries very prominent, 
very often nearly white and extending from costal to inner margin (Figs. 2, 4, 6 & 
8). Nominate appalachia females may in some individuals show markedly lighter 
subapical areas above and lighter limbal areas below than female figured though 
never as in leeuwi females. 

Holotype male: Wakelee, Cass Co., Michigan, 24 July 1972, leg. Irwin Leeuw; 
deposited temporarily in the senior author’s collection. 

Allotype female: Wakelee, Cass Co., Michigan, 4 July 1957, leg. M. C. Nielsen; 
deposited in the collection of Michigan State University. 

Paratypes: Michigan. Cass County: 2 males, 30 June 1972; 2 males, 24 July 
1972; 3 males, 3 July 1973 (leg. Irwin Leeuw); 1 male, 16 July 1967; 1 female, 16 
July 1970; 6 males, 15 July 1972 (leg. P. J. Conway); 1 male and 1 female, 7 July 
1971; 2 males, 9 July 1971 (leg. R. R. Irwin); 1 male (abdomen missing), 1 July 
1973 (leg. M. G. Seaborg). Clinton County: 4 males, 16 July 1972 (leg. M. C. 
Nielsen). Lenawee County: 1 male, 20 June 1970; 1 male, 15 July 1973; 1 female, 
22 July 1973 (leg. M. C. Nielsen). St. Joseph County: 1 male, 12 July 1972 (leg. 
M. C. Nielsen). Barry County: 1 male, 12 July 1956 (leg. R. L. Fischer); 1 male, 
14 July 1973 (leg. M. C. Nielsen). Montcalm County: 1 male, 3 July 1952; 1 male, 
9 July 1953 (leg. M. C. Nielsen). Wayne County: 1 male, no date (leg. A. W. 
Andrews); 1 female, 10 July 1943 (leg. M. C. Nielsen). Washtenaw County: 1 
female, 18 July 1964 (collector unknown). 

The 31 males and 6 females of the type series are deposited in the 
following collections: Michigan State University, Illinois Natural History 
Survey, P. J. Conway, M. C. Nielsen, R. R. Gatrelle, R. T. Arbogast, 
Irwin Leeuw and M. G. Seaborg. 

In addition to the type series, material referable to leewwi was ex- 
amined from Devil's Lake State Park, Wisconsin; Illinois Beach State 
Park, Lake County, Illinois; Paulding and Lake Counties in Ohio; Bed- 
ford and Reading, Pennsylvania; and Martha’s Vineyard, Massachusetts. 
Five males from Baltimore, Maryland which we examined were darker 
than typical leeuwwi, but were still closer to the new subspecies than to 
nominate appalachia. 

Eighty specimens were examined in the course of this study of nomi- 
nate southern appalachia. We were not able to examine specimens of 
L. appalachia appalachia from the type locality, Connestee Falls, near 


Figs. 9-14. Lethe appalachia leeuwi (Gatrelle and Arbogast), new subspecies: 
9 & 12, paratype male, Montcalm County, Michigan, 3 July 1952; 10 & 13, paratype 
male, Lenawee County, Michigan, 20 June 1970; 11 & 14, female, Paulding County, 

Ohio, 11 July 1971. These three specimens show the variation within the new sub- 

VoLUME 28, NUMBER 4 363 

Brevard, North Carolina, but we did examine material from Table Rock 
State Park, Pickens County, South Carolina which is just 13 miles south 
of Brevard. We found these specimens to agree in every respect with 
Chermock’s description of appalachia. A note of interest here is that the 
type locality, Connestee Falls area, is now undergoing drastic change. 
The area is being turned into a “resort” community and housing develop- 
ment. The drastic changes in the environment may well lead to the 
extinction of appalachia in that area. The specimens from Table Rock, 
South Carolina are of the same phenotype as those from Brevard and 
unless a colony is located closer than 13 miles to the type locality, the 
Table Rock populations may be the closest thing to topotypes available 
to the taxonomist. The specimens which we examined of nominate 
appalachia were from the following areas. South Carolina: Pickens, 
Dorchester and Charleston Counties. Georgia: Fannin and Cherokee 
Counties, and the Atlanta area. Florida: Pasco County. Mississippi: 
Pontotoc, Lee, Lafayette, Choctaw, Oktibbeha, and Winston Counties. 
All the specimens from these localities closely resembled the material 
from Table Rock except that the specimens from Pasco County, Florida 
averaged somewhat darker. 

Our thanks go to the many persons who loaned us material for examina- 
tion and who helped us with their ideas. 


Carpe, R. T., A. M. SHaprro & H. K. CLencu. 1970. Sibling species in the eury- 
dice group of Lethe (Lepidoptera; Satyridae). Psyche 77: 70-103. 

CuHEerMock, R. L. 1947. Notes on North American Enodias (Lepidoptera). En- 
tomol. News. 58: 29-35. 

Brown, L. N. 1973. A population of Lethe appalachia (Satyridae) from West 
Central Florida. J. Lepid. Soc. 27: 238-239. 


The Charles Rudkin collection of Lepidoptera has been acquired by the Museum 
of Systematic Biology, University of California, Irvine. The collection contains over 
10,000 mounted specimens (in modified Riker Mounts), primarily Rhopalocera. The 
collection is especially rich in California material, but also contains a fair amount of 
material from southeastern Arizona and the South Pacific. Nearly all specimens were 
collected from 1930-1945. Rudkin’s field notebooks and other memorabilia will 
accompany the collection. 

Larry J. Orsax, Museum of Systematic Biology, School of Biological Sciences, 
University of California, Irvine, California 92664. 



Dominick et al., editors. E. W. Classey Ltd., and R. B. D. Publications Inc. _ Dis- 
tributed in North America by Entomological Reprint Specialists, P.O. Box 77971, 
Dockweiler Station, Los Angeles, California 90007. 

One of the greatest needs of students of the Lepidoptera of North America has 
been a definitive work on our moths, which number in excess of 10,000 species. It 
is practically impossible to accurately name many of the species, with the exception 
of some genera and a few higher groups that have been recently studied; these re- 
visionary studies are often scattered in the literature and are not necessarily easily 
available to all collectors. Now this need is being filled most adequately with the 
series of definitive studies that will make up this series. A total of over 50 fascicles 
are planned, with three or four to be published each year; hopefully the task will 
be completed in the next 12 years or so. 

Each fascicle may cover several small families, one family, or a part of a large 
family, and in itself, is a taxonomic revision of the group being covered. New taxa 
of all ranks are described where needed; old ones are redescribed. Keys are pro- 
vided to help in identification. Each species account includes a reference to the 
original description and to synonyms when present. Each species is diagnosed, its 
variation discussed, and its distribution, habitat, and whatever is known about its 
life history and foodplants given. Genitalia are described, and figured when pertinent. 
In addition, each species, its subspecies and color variants are illustrated in full color 
and natural size; the smaller species are being enlarged. 

A standard format is being used throughout the series. The one exception is in 
the usage of subspecific names. The board of directors, including the four authors 
that have published fascicles to date, could not agree on this problem; Ferguson and 
Munroe use this concept, while Franclemont and Hodges do not. Ferguson and 
Franclemont, in their respective fascicles reviewed below, outline the pros and cons 
of the question. Unfortunately, Hodge’s work appeared before this subject was 
clarified; the reader, not realizing this, may be puzzled by the way the taxa are 

Each fascicle is a sumptuous example of printers’ art. Every one is of large size, 
beautifully printed on excellent paper, and contains some of the best color plates 
ever printed of our North American moths. To produce all this entails great expense; 
consequently the price per fascicle is relatively high. However, considering the 
above factors, plus the fact that this series will be the standard source of reference 
for generations to come, I know that this is money well spent. (In some ways, the 
Moths of North America is comparable to the Biologia Centrali-America and Seitz’ 
Macrolepidoptera of the World; have you tried pricing or even finding copies of 
these to buy recently? ) 

This publication is being called the definitive work on our North American moths, 
and I try to judge the individual fascicles according to this simple definition. To 
achieve this status each author should know the group thoroughly, not only in North 
America but in other parts of the world, be thoroughly knowledgeable about the 
pertinent literature, have examined the types of valid names and synonyms (and 
designating lectotypes where needed), and studied the bulk of specimens in this 
country. It is also preferable to have authors that have spent considerable time, 
over the years, working with the group prior to publishing on it; this has not always 
been possible—there just aren’t enough competent specialists to properly cover each 
and every group of the moths. 

VoLuME 28, NUMBER 4 365 

Fascicle 21. SpHiNncomEA: SPHINGIDAE, by Ronald W. Hodges. 1971. xii + 158 p., 
16 pls. (14 in color), 8 halftones, 19 text figs. 

The text on the hawkmoths is a thoroughly competent piece of popular writing 
on this group of mostly large-sized moths, numbering 115 species in the area covered. 
No one should have very much trouble determining the different species in this 
family. Hodges does not believe in the subspecies concept; unfortunately he did 
not mention this matter in the introduction to his paper. This will cause some con- 
fusion; geographic variation is discussed within the different species, but the names 
that have heretofore been used in the subspecific sense are merely listed in the 
species synonymies. 

Hodges uses as a higher classification one apparently modified from Carcasson, 
thus differing from both McDunnough’s 1938 Check List and Forbes’ 1948 Lepi- 
doptera of New York; the latter two were based primarily on Rothschild and Jordan 
(1903). For many readers it may be the first time they have been introduced to 
this new higher classification; it would have been helpful if Hodges had gone into 
greater detail comparing the two, amplifying the reasons for this change. 

The listing of the supraspecific categories is followed by a key to the genera; 
Hodges does not have a key for the subfamilies or tribes, although they are defined 
in the text. This, in turn, is followed by “partial” keys to the genera based on the 
pupae (after Mosher, 1918) and on the mature larvae (after Forbes, 1911). It is 
possible that both the latter could have been modernized with relatively little effort. 

In my opinion, this fascicle does not attain the status of a “definitive work.” If 
Hodges had taken the time to visit the American Museum of Natural History, for 
instance, he would have added one or two more species to the work; additional 
distributional data and information on flight periods would have been added for at 
least 30 species—almost one-fourth of the number covered in his work. There are 
some two dozen errors in the bibliographical citations, either in the references them- 
selves or incorrectly giving the original combination for the names. At least two 
references to early stages and foodplants were overlooked. 

Notwithstanding this list of criticisms, the paper is the best one ever to appear 
on our North American sphingids, and I would strongly advise anyone interested in 
the fascinating group to obtain a copy. 

Fascicle 20.2. BomMBycomDEA: SATURNIIDAE, by Douglas C. Ferguson. Part 2A, 
Citheroniinae, Hemileucinae (in part); 1971, p. 1-153, pls. 1-11 (color), text figs. 
1-19. Part 2B, Hemileucinae (in part), Saturniinae; 1972, p. 154-275 + xxi, pls. 
12-22 (color), text figs. 20-30. 

This large family is basically tropical and subtropical in distribution, with about 
65 species occurring in America north of Mexico. Ferguson did a much more 
thorough job of research and study than did Hodges; this is quickly recognizable 
when one reads the text. The last is excellently done; in fact, in my opinion it is one 
of the best written and most complete studies I have had the pleasure of reading, 
as Ferguson did an excellent job in combining the popular and scientific aspects of 
the subject. Of particular value is the effort that was made to fully explain many 
of the “sticky” problems in this group; it is such attention to detail that increases 
the value of this publication. 

Ferguson basically follows the suprageneric classification of Michener (1952), 
deviating mainly in raising the appropriate subgenera to full generic status in most 
cases. Keys are presented to the subfamilies based on the adults, last instar larvae, 
and pupae (after Mosher, 1916); within each subfamily there are similar keys to 
the included genera. Special emphasis is given to life histories, foodplants, and the 
morphological characters of the larvae and pupae. 

The bibliographical references and text are relatively free of mistakes and omissions. 
This fascicle indeed lives up to the advance billing of a definitive work. 


Fascicle 13.1. PyRALOIDEA (IN PART), by Eugene Munroe. Part 1A, Scopariinae, 
Nymphulinae; 1972, p. 1-134. Part 1B, Odontiinae, Glaphyriinae; 1972, p. 135-250. 
Part 1C. Evergestinae; “1973” [1974], p. 251-304 + xx, pls. 1-13 (color), A-K 
(halftones ). 

Munroe has spent more than 25 years studying the Pyraloidea in general and the 
North American fauna in particular. He has done field work in practically every 
part of continental North America, and has visited most of the major collections and 
institutions in this area. His studies have taken him throughout Europe and to their 
museums, to Africa and to tropical America. He has built up an encyclopedic 
knowledge of the world fauna of the pyralids and, based on this, is setting forth the 
first comprehensive manual for the identification of the North American species of 
all families and subfamilies of the Pyraloidea. The system of classification he is 
proposing for our fauna is considerably different from what we have had before, 
with the introduction of subfamilies and tribes that are new to us. Four families 
are involved; one of these, the Pyralidae, is divided into 16 subfamilies. Part 1A 
contains the definition of the superfamily and of the Pyralidae; keys are provided 
to separate the families and subfamilies of the previously-mentioned family. Five 
of the subfamilies are covered in fascicle 13.1; each group has keys to the genera and 
species (when more than one is known). A number of tribes, genera, species, and 
subspecies are described as new. In a few cases Munroe may be splitting the taxa 
a bit too thin. Some are admittedly, “perhaps not really worth separating,’ as in 
some of the genera of Evergestinae, where the only differences are in the variously- 
shaped frontal prominences on the head. Similarly, he recognizes as distinct some 
species that are apparently morphologically indistinguishable except for differences 
in wing color only. 

As in the other parts of this series, each species and subspecies is illustrated on 
the color plates. For the great majority, it is the first time they have been figured. 
The quality of some of these color plates does not seem to me to be quite as good 
as those in the other fascicles, but this is undoubtedly due to the smaller size of 
the moths, more specimens per plate (with some distracting pins being shown for 
the first time), and the magnification. Nevertheless, these color plates continue to 
be the best ever produced of our North American moths. 

Parts 1A and 1B each have a modest number of mistakes in the bibliographical 
references; 1C is greatly improved, as I did not note a single one. Munroe is to be 
congratulated for designating lectotypes where necessary. I would like to see the 
work on types extended one additional step, with the depository of each being desig- 
nated. This should not appreciably increase the length of the reference section of 
each name, and it would be an invaluable aid to present and future workers. 

Munroe is to be congratulated for a piece of original work excellently done; I am 
eagerly looking forward to succeeding fascicles in his monumental work on our 

LODIDAE, BOMBYCIDAE, LASIOCAMPIDAE, by John G. Franclemont. 1973. viii + 86 p., 
11 color pls., 22 text figs. 

Franclemont erects the superfamily Mimallonoidea for the single anomalous family 
Mimallonidae (Lacosomidae of McDunnough’s Check List, 1938). The four included 
species are placed in three genera; of these, one genus and one species are described 
as new. Three families are included in this section of the Bombycoidea; all are 
small-sized in our area. The Apatelodidae (Zanolidae of McDunnough, 1938) con- 
tains two genera and five species; the Bombycidae contains only the introduced silk- 
worm, Bombyx mori (Linnaeus); and the Lasiocampidae encompass 12 genera with 
about three times that number of species. 

The main contribution in this fascicle is a new suprageneric classification of the 

VoLUME 28, NUMBER 4 367 

Lasiocampidae, which is divided into three subfamilies and one of these into two 
tribes. The Neotropical representatives of this family are relatively unknown and 
much work needs to be done with them; when properly studied it will be interesting 
to find out how these species and genera fit into Franclemont’s classification. 

There are only two relatively large genera in our Lasiocampidae. One of these, 
Malacosoma, has recently been revised by Stehr & Cook (1968); this work is closely 
followed in the fascicle, and their key to the mature larvae, as well as pls. 1 and 2 
of larvae, are taken directly. Throughout the discussion of this genus Franclemont 
continually refers to “the revision of the American species by Stehr (1968).” While 
it is true that Stehr did most, or all, of the work in that revision, it is assumed that 
Franclemont worked from the published revision; if this is so, then the correct 
reference should read Stehr & Cook, 1968, as this is how the results were published. 

The other large genus is Tolype, and this represents the main original research on 
a specific level by Franclemont in the fascicle. He admits, in the Introduction, that 
there are a number of problems here, particularly in western North America, and 
that he did not have the time to try to solve them properly. Independently, and 
prior to the publication of this fascicle, I had studied our collection of this genus, 
mainly utilizing genitalic dissections. Franclemont’s treatment of the few eastern 
species appears sound; in the western part of the continent his handling of the species 
leaves quite a bit to be desired — he, indeed, did not solve all the problems. One 
thing that might have helped him would have been to study the extensive material 
at the American Museum of Natural History. I cannot help but get the feeling that 
Franclemont has only hazy notions about distributional patterns of western species. 
For example, he gives the distribution of Tolype dayi Blackmore (on p. 44) as 
British Columbia, Washington, and Montana; yet, on pl. 3, fig. 34, he illustrates a 
specimen of dayi from Santa Cruz Co., California, which is a good 700 miles away. 
I had trouble comparing the drawings of the female genitalia, particularly those of 
the sterigma, with my dissections. Franclemont may have completely overlooked 
one character in the female, as nowhere did I find reference to the nature of the 
scales in the anal tuft; these may have good specific characters. 

One point that surprised me was Franclemont’s apparent ignorance of the litera- 
ture. Three examples: he gave incorrect designations for the type species of both 
Tolype and Artace; for Tolype dayi he states that “the larva has not been described,” 
but in reality the description has appeared twice. There are at least five other 
bibliographical errors. 

The subspecies problem is admirably handled in this fascicle. Franclemont gives 
his views on why he does not utilize this concept in the Introduction. In the dis- 
cussion of variability within the individual species he clearly points out when geo- 
graphic variation occurs and if a name is available for that population. However, 
he insists on continually using the unmodified word “race” in place of subspecies or 
geographic subspecies. His term is not recognized by the Code; it always strikes 
me that this is a quaint Victorian term that is more suitable for the Olympics than 
for inclusion in a major entomological systematic work being published today. 

Over-all, this fascicle has many excellent points. However, it also has a surprising 
number of drawbacks for the coverage of such a small group that entailed relatively 
little original research. I can hardly consider this as a definitive piece of work. 

FreDERICK H. Rinpce, Department of Entomology, the American Museum of 
Natural History, Central Park West at 79th Street, New York, New York 10024. 



ALEX K. WYATT (1878-1971) 

An entomological career of three-quarters of a century was brought to 
a close with the death of Alex K. Wyatt in Chicago, [linois on May 14, 
1971. At first known to the entomological world as Alexander Kwiat, he 
changed his name to its present form in 1918. He was born in Chicago, 
December 28, 1878, the son of German immigrant parents. He married 
Eva Stuhlfaut in September, 1908 and to them were born three children: 
Elva A. (Mrs. J. T. Mauer) of Chicago, Lillian M. (Mrs. Leslie Skutle) 
of Kent, Ohio, and Harold, who died in April, 1930 at the age of ten. 

Mr. Wyatt was educated in the public schools of Chicago. He grad- 
uated from Newberry grammar school in 1892 and for two years at- 
tended North Division High School, whose principal was Oliver S. 
Westcott, himself an entomologist. Following eight months’ attendance 
at business college he secured a position as office boy with a real estate 
firm in 1895. Except for two years in the office of a fire insurance com- 
pany, the rest of his business career was spent in the real estate field. 
He operated his own industrial real estate brokerage business, from which 
he retired in 1956. 

His interest in Lepidoptera arose at an early age, and developed during 
his second year in high school, when he learned collecting and preserving 
techniques under the tutelage of Westcott. Beginning with a general 

VOLUME 28, NUMBER 4 369 

insect collection, he soon found this to be too great an undertaking, and 
disposed of all his specimens except Lepidoptera by giving them to 
C. T. Brues and A. L. Melander, who were also pupils of Westcott and 
who later became prominent professional entomologists. His earliest 
collecting was done in Chicago’s Lincoln Park before butterfly nets were 
prohibited there. He soon became acquainted with John L. Healy, Arthur 
J. Snyder, W. E. Longley, James Tough and others who were associated 
with the Chicago Academy of Sciences. This group organized itself in 
1897 as the Chicago Entomological Society, which Wyatt served as sec- 
retary during most of its existence. He was also a charter member of 
the Lepidopterists’ Society and of the Entomological Society of America. 

Wyatt was the last survivor of Chicago’s fraternity of Bohemian col- 
lectors, which included Paul Vollbrecht, Berthold Neubarth, Charles 
Krueger, Arthur Herz and others, most of whom belonged to a German 
social club with headquarters near Lincoln Avenue and Belmont Street. 
Its members made regular collecting trips to such favorite Chicago area 
localities as Palos Park and Schiller Park, Illinois and Hessville (now 
part of Hammond), Indiana. Also among his close friends were Murray 
O. Glenn, John G. Franclemont, and the late Otto Buchholz, Emil Beer 
and Henry Ramstadt. Most of Wyatt’s collecting was done near Chicago, 
but he made collecting trips at various times to Oregon and Washington, 
Kentucky and Tennessee, and several to Florida. 

Following his retirement from business, Wyatt in 1957 donated his 
collection to the Field Museum of Natural History, Chicago. It consisted 
of 24,644 specimens including about 5000 species and varieties, as well 
as holotypes of taxa he described and an undetermined number of para- 
types. 2295 specimens representing some 500 species were butterflies. 
At the same time he joined the Museum staff as a research associate in 
the Division of Insects. He personally incorporated his collection into 
that of the Museum, while supervising a general rearrangement of the 

In 1959 he became afflicted with heart disease and cataracts on both 
eyes. He and his wife spent the winter of 1959-60 in St. Petersburg, 
Florida. There he collected at store fronts almost every evening, securing 
a total of more than 2500 specimens during six months, all of which were 
deposited in the Museum. In the summer of 1960 he underwent surgery 
for the cataracts, and although the operation itself was successful, retinal 
complications followed and his vision deteriorated to the extent that he 
could no longer drive a car nor determine specimens, which brought to 
an end his collecting activities. Following the death of his wife in 
November, 1962 he made his home with his daughter, Mrs. Mauer. De- 


spite his advanced age and physical handicaps he continued to visit the 
Museum fairly regularly for several more years. 

Wyatt was particularly interested in Holomelina and Papaipema, 
Heliothiinae, and in life history research and the collecting and rearing 
of larvae. His many contributions to the knowledge of the life history 
of moths are found in the literature under his own authorship as well as 
that of others. He was adept at fashioning his own equipment, much of 
which continues in use today. One of his outstanding characteristics was 
an unfailing willingness to aid and encourage younger lepidopterists, 
among them this author. 

Lepidopterous taxa named in Wyatt's honor include Lycaena thoe ab. 
wyatti Gunder, Lasionycta wyatti Barnes and Benjamin, Papaipema in- 
quaestia form wyatti Barnes and Benjamin, and Eteobalea wyattella 
(Barnes and Busck). 

The author acknowledges with sincere appreciation the cooperation of 
Mrs. Elva Mauer, Mr. Murray O. Glenn, and especially Mr. Henry Dybas, 
Curator of Insects at the Field Museum, in the preparation of this article. 
The accompanying photograph, taken in 1961, was provided through 
the courtesy of that institution. Portions of the article were adapted from 
unpublished autobiographical material of Wyatt in the museum’s archives. 


In addition to the papers listed below, Wyatt was the author of minutes 
of the Entomological Section of the Chicago Academy of Sciences, which 
were published from time to time in the Entomological News. His 
earliest papers appeared under the name of Alexander Kwiat. 

1908. One day’s collecting, with a description of a new noctuid. Entomol. News 
19; 420-424 

1916. Collecting Papaipemae (Lepidoptera). Entomol. News 27: 228-234. 

1926. [John L. Healy]. Entomol. News 37: 128. 

1927a. Collect